Panthera spelaea: Difference between revisions

Panthera spelaea; Temporal range: Middle-Late Pleistocene, 0.6–0.013 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
	Skeleton in Natural History Museum, Vienna
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Suborder:	Feliformia
Family:	Felidae
Subfamily:	Pantherinae
Genus:	Panthera
Species:	†P. spelaea
Binomial name
	†Panthera spelaea; Goldfuss, 1810
Subspecies
	Panthera spelaea vereshchagini (Baryshnikov & Boeskorov, 2001); Panthera spelaea spelaea Goldfuss, 1810;
	Red indicates the maximal range of Panthera spelaea, blue Panthera atrox, and green Panthera leo.
Synonyms
	Panthera leo spelaea M. Boule & L. De Villeneuve, 1927;

Browse history interactively

← Previous edit

Content deleted Content added

VisualWikitext

Inline

Latest revision as of 17:34, 17 November 2024

Panthera spelaea, commonly known as the cave lion (or less commonly as the steppe lion), is an extinct Panthera species that was native to Eurasia and northwest North America during the Pleistocene epoch. Genetic analysis of ancient DNA has revealed that while closely related, it was a distinct species genetically isolated from the modern lion (Panthera leo),^[1] with the genetic divergence between the two species estimated at around 500,000 years ago.^[2] The earliest fossils of the P. spelaea lineage (either regarded as the separate species Panthera fossilis or the subspecies P. spelaea fossilis) in Eurasia date to around 700,000 years ago (with possible late Early Pleistocene records).^[3] It is closely related and probably ancestral to the American lion (Panthera atrox).^[2] The species ranged from Western Europe to eastern Beringia in North America, and was a prominent member of the mammoth steppe fauna, and an important apex predator across its range. It became extinct about 13,000 years ago.^[4] It closely resembled living lions with a coat of yellowish-grey fur though unlike extant lions, males appear to have lacked manes.

Panthera spelaea interacted with both Neanderthals and modern humans, who used their pelts and in the case of the latter, depicted them in artistic works.

Taxonomy

Felis spelaea was the scientific name used by Georg August Goldfuss in 1810 for a fossil lion skull that was excavated in a cave in southern Germany.^[5] It possibly dates to the Würm glaciation.^[6]^[1]^[4]

Several authors regarded Panthera spelaea as a subspecies of the modern lion, and therefore as Panthera leo spelaea.^[7]^[8]^[9]^[1] One author considered the cave lion to be more closely related to the tiger based on a comparison of skull shapes, and proposed the scientific name Panthera tigris spelaea.^[10]

Results from morphological studies showed that it is distinct in cranial and dental anatomy to justify the specific status of Panthera spelaea.^[11]^[12] Results of phylogenetic studies also support this assessment.^[13]^[14]^[15]

In 2001, the subspecies Panthera spelaea vereshchagini was proposed for seven specimens found in Siberia and Yukon, which have smaller skulls and teeth than the average P. spelaea.^[16] Before 2020, genetic analysis using ancient DNA provided no evidence for their distinct subspecific status; DNA signatures from P. spelaea from Europe and Alaska were indistinguishable, suggesting one large panmictic population.^[14]^[17] However, analysis of mitochondrial genome sequences from 31 cave lions showed that they fall into two monophyletic clades. One lived across western Europe and the other was restricted to Beringia during the Pleistocene. For this reason, the Beringian population is considered a distinct subspecies, P. s. vereshchagini.^[17]^[18]

Evolution

Lion-like pantherine felids first appeared in the Tanzanian Olduvai Gorge about 1.7 to 1.2 million years ago. These cats dispersed into Eurasia from East Africa around the end of the Early Pleistocene and the beginning of the Middle Pleistocene, giving rise to Panthera fossilis. The oldest widely accepted fossils of P. fossilis in Europe date to around 700,000 years ago,^[19]^[20]^[3]^[21] with possible older fossils from Western Siberia dating to the late Early Pleistocene.^[22] Different authors considered Panthera fossils as either a distinct species ancestral to P. spelaea,^[23] or as a subspecies of P. spelaea.^[21]^[24] Recent nuclear genomic evidence suggest that interbreeding between modern lions and all Eurasian fossil lions took place up until 500,000 years ago, but by 470,000 years ago, no subsequent interbreeding between the two lineages occurred.^[19]^[1]^[2]

The following cladogram shows the genetic relationship between P. spelaea and other pantherine cats.^[15]

Felis catus

Pantherinae

Neofelis nebulosa (clouded leopard)

Panthera

Panthera tigris (tiger)

Panthera onca (jaguar)

Panthera uncia (snow leopard)

Panthera pardus (leopard)

Panthera leo

	Panthera leo leo (Asian, North and West African lions)

	Panthera leo melanochaita (Southern and East African lions)

†Panthera spelaea

Specimens intermediate between P. fossilis and Late Pleistocene P. spelaea are referred to as the subspecies P. s. intermedia.^[20] The transition from P. fossilis to Late Pleistocene P. spelaea involved significant changes in skull and tooth morphology.^[25] Mitochondrial DNA sequence data from fossil lion remains show that the American lion represents a sister group of Late Pleistocene P. spelaea, and likely arose when an early P. spelaea population became isolated south of the Cordilleran Ice Sheet. Initially this was suggested to be around 340,000 years ago,^[14] but later studies suggested that the split between the two species was probably younger, around 165,000 years ago, consistent with the late first appearance of P. spelaea in Eastern Beringia (now Alaska and adjacent regions) during the Illinoian (around 190-130,000 years ago).^[26]

Characteristics

Carvings and cave paintings of cave lions, which were discovered in the Lascaux and Chauvet Caves in France, were dated to 15,000 to 17,000 years old.^[27]^[28] A drawing in the Chauvet cave depicts two cave lions walking together. The one in the foreground is slightly smaller than the one in the background, which has been drawn with a scrotum and without a mane.^[29] Such cave paintings suggest that male cave lions completely lacked manes, or at most had very small manes.^[4]

Early members of the cave lion lineage (including those assigned Panthera fossilis) during the Middle Pleistocene were considerably larger than individuals of P. spelaea from the Last Glacial Period and modern lions, with some of these individuals having an estimated length of 2.5–2.9 metres (8.2–9.5 ft), shoulder height of 1.4–1.5 metres (4.6–4.9 ft) and body mass of 400–500 kilograms (880–1,100 lb), respectively, making them among the largest cats to have ever lived. The Late Pleistocene Panthera spelaea spelaea was noticeably smaller though still large relative to living cats, with an estimated length and shoulder height of 2–2.1 metres (6.6–6.9 ft) and 1.1–1.2 metres (3.6–3.9 ft), respectively, The species showed a progressive size reduction over the course of the Last Glacial Period up until its extinction, with the last P. spelaea populations comparable in size to small-sized modern lions, with a body mass of only 70–90 kilograms (150–200 lb) and a body length and shoulder height of only 1.2–1.3 metres (3.9–4.3 ft) and 70–75 centimetres (2.30–2.46 ft) respectively.^[24]^[30]

P. spelaea had a relatively longer and narrower muzzle compared to that of the extant lion. Despite this, the two species do not exhibit major differences in morphology.^[4] Like modern lions, females were smaller than males.^[31]

In 2016, hair found near the Maly Anyuy River was identified as cave lion hair through DNA analysis. Comparison with hair of a modern lion revealed that cave lion hair was probably similar in colour as that of the modern lion, though slightly lighter. In addition, the cave lion is thought to have had a very thick and dense undercoat comprising closed and compressed yellowish-to-white wavy downy hair with a smaller mass of darker-coloured guard hairs, possibly an adaptation to the Ice Age climate.^[32] While juveniles fur coat colour was yellowish, adult cave lions are suggested to have had grey fur.^[18]

Distribution and habitat

During the Last Glacial Period, P. spelaea formed a contiguous population across the mammoth steppe, from Western Europe to northwest North America.^[4]^[33] It was widely distributed in the Iberian Peninsula,^[34] Italian Peninsula,^[35] Southeast Europe,^[36] Great Britain,^[4] Central Europe,^[37]^[38]^[39] the East European Plain,^[4] the Ural Mountains,^[40] most of Northeast Asia (ranging as far south as Northeast China^[41] and possibly the Korean peninsula^[42]), and across the Bering land bridge into Alaska and Yukon.^[4] The cave lion had a wide elevation range, with finds extending up over 2,000 metres (6,600 ft) above sea level in the European Alps and in Buryatia in Northern Asia, though they probably did not occupy mountainous habitats all-year round.^[43] The cave lion probably inhabited predominantly open habitats such as steppe and grasslands although it would have also have occurred in open woodlands as well.^[4] It may have sought out hibernating bears in montane caves as a food source during the winter.^[44] While during the Last Glacial Period it was often associated with cold environments, the species also inhabited temperate environments,^[45] such as in Europe during the Last Interglacial/Eemian.^[46]

Paleobiology

Ecology

P. spelaea was one of the keystone species of the mammoth steppe, being one of the main apex predators alongside the gray wolf, cave hyena and brown bear.^[48] Large amounts of bones belonging to P. spelaea were excavated in caves, where bones of cave hyena, cave bear and Paleolithic artefacts were also found.^[49]^[50] Despite their common name, "cave lions" probably only infrequently if ever used caves, and were present in regions where caves were absent.^[51] Some of these accumulations of cave lion bones in cave hyena dens have been attributed to confrontations between cave hyenas and cave lions over carcasses, with the remains of cave lions killed in these confrontations subsequently transported to the dens.^[52]^[51]

Isotopic analyses of bone collagen samples extracted from remains in Europe^[53]^[48] and East Beringia^[54] indicate that reindeer were particularly prominent in the diet of cave lions in these regions during the Last Glacial Period.^[48] Cave lions also seem to have opportunistically preyed on the cubs of cave bears.^[48]^[53] Isotopic analysis of other European specimens suggests a diet including wild horse, woolly mammoth and cave bears for these individuals.^[48]

Other possible prey species were giant deer, red deer, muskox, aurochs, wisent, steppe bison, and young woolly rhinoceros. It likely competed for prey with the European leopards, cave hyenas, brown bears and grey wolves in Eurasia,^[55] along with short-faced bears, Homotherium, and Beringian wolves in Beringia.^[48]

Social behavior

Whether or not cave lions existed in prides like modern lions is unclear. Isotopic analysis done by Hervé Bocherns in 2015, suggested cave lions in Europe may have been solitary, due to scattering of individual data which was more similar to individualistic behavior compared to modern day lion populations.^[48] Some other authors have also argued that the absence of manes in cave lions suggests that cave lions did not live in prides, given the importance of manes in the social hierarchy of modern lions.^[18] Boeskorov et al. 2021 suggested both European and Beringian cave lions may have hunted in larger prides than modern lions because sexual dimorphism in cave lions was more pronounced than in modern African lions and solitary big cats. However, they admitted the data is insufficient to come down to a certain conclusion.^[18]

Cave lion cubs appear to have lived in dens during their earliest stages of life, like modern lion cubs and were likely solely raised by females, like living Panthera species.^[18]

Relationship with humans

Drawing of a now-lost Palaeolithic sculpture of a cave lion from Isturitz, France

Drawing of cave lions in the Chamber of Felines, Lascaux caves

Carving of a Panthera spelaea head from Vogelherd Cave, dating to around 35–40,000 years ago

Anthropomorphic Lion-man (Löwenmensch) figurine, around 35-41,000 years old

Cave lions were hunted and their pelts exploited in Europe by Neanderthals during the Middle Paleolithic,^[56] and during the Upper Paleolithic by modern humans in Spain as evidenced in the La Garma site dating to the Magdalenian.^[57] Modern humans also drew cave paintings of cave lions, engraved their likeness on bones and created sculptures of them, including the famous anthropomorphic lion-man figure from Hohlenstein-Stadel cave in Germany dating to around 41-35,000 years ago with the body of a human and the head of a lion. Cave lion canines with perforated holes may have been worn as personal ornaments.^[56] Decorated stones with engravings representing cave lions have been found in southern Italy.^[58]

Extinction

Radiocarbon dating suggests that the species went extinct approximately simultaneously across its range during the last few thousand years of the Late Pleistocene, around 14-15,000 years ago, possibly surviving around 1000 years later in the far east North American portion of its range. This timing roughly corresponds to the onset of the Bølling–Allerød Interstadial warm period and the consequent collapse of the mammoth steppe ecosystem. The precise cause of its extinction is unclear, but possibly involved environmental change from open habitats to closed forests, changes in prey abundance, as well as human impact, though it is difficult to distentangle the precise causes of its extinction.^[4] Cave lions appear to have undergone a population bottleneck that considerably reduced their genetic diversity between 47,000 and 18,000 years ago, probably driven at least in part by climatic instability.^[59]

Mummified specimens

In 2008, a well-preserved mature cave lion specimen was unearthed near the Maly Anyuy River in Chukotka Autonomous Okrug, which still retained some clumps of hair.^[60]

In 2015, two frozen cave lion cubs, estimated to be between 25,000 and 55,000 years old, were discovered close to the Uyandina River in Yakutia, Siberia in permafrost.^[61]^[62]^[63] Research results indicate that the cubs were likely barely a week old at the time of their deaths, as their milk teeth had not fully erupted. Further evidence suggests the cubs were hidden at a den site until they were strong enough to follow their mother back to the pride, as with modern lions. Researchers believe that the cubs were trapped and killed by a landslide, and that the absence of oxygen underground hindered their decomposition and allowed the cubs to be preserved in such good condition. A second expedition to the site where the cubs were found was planned for 2016, in hopes of finding either the remains of a third cub or possibly the cubs' mother.^[64]

In 2017, another frozen specimen, thought to be a lion cub, was found in Yakutia on the banks of the Tirekhtyakh River (Russian: Тирехтях), a tributary of the Indigirka River. This male cub was thought to be slightly older than the 2015 cubs at the time of its death; it is estimated to have been around one and a half to two months.^[65] In 2018, another preserved carcass of a cub was found in a location 15 m (50 ft) away. It was considered to be around a month old when it died approximately 50,000 years ago, and presumed to be a sibling of the male cub.^[66] However, carbon dating showed them to have lived about 15,000 years apart, with the female estimated to have lived 28,000 years ago, and the male 43,448 years ago.^[18] Both cubs were well preserved, albeit with a few damages, with the female possibly being the "best preserved" animal discovered from the Ice age.^[67]

References

^ ^a ^b ^c ^d Burger, J.; Rosendahl, W.; Loreille, O.; Hemmer, H.; Eriksson, T.; Götherström, A.; Hiller, J.; Collins, M. J.; Wess, T.; Alt, K. W. (2004). "Molecular phylogeny of the extinct cave lion Panthera leo spelaea". Molecular Phylogenetics and Evolution. 30 (3): 841–849. Bibcode:2004MolPE..30..841B. doi:10.1016/j.ympev.2003.07.020. PMID 15012963.
^ ^a ^b ^c Manuel, M. d.; Ross, B.; Sandoval-Velasco, M.; Yamaguchi, N.; Vieira, F. G.; Mendoza, M. L. Z.; Liu, S.; Martin, M. D.; Sinding, M.-H. S.; Mak, S. S. T.; Carøe, C.; Liu, S.; Guo, C.; Zheng, J.; Zazula, G.; Baryshnikov, G.; Eizirik, E.; Koepfli, K.-P.; Johnson, W. E.; Antunes, A.; Sicheritz-Ponten, T.; Gopalakrishnan, S.; Larson, G.; Yang, H.; O'Brien, S. J.; Hansen, A. J.; Zhang, G.; Marques-Bonet, T. & Gilbert, M. T. P. (2020). "The evolutionary history of extinct and living lions". Proceedings of the National Academy of Sciences of the United States of America. 117 (20): 10927–10934. Bibcode:2020PNAS..11710927D. doi:10.1073/pnas.1919423117. PMC 7245068. PMID 32366643.
^ ^a ^b Marciszak, Adrian; Schouwenburg, Charles; Gornig, Wiktoria; Lipecki, Grzegorz; Mackiewicz, Paweł (November 2019). "Morphometric comparison of Panthera spelaea (Goldfuss, 1810) from Poland with the lion remains from Eurasia over the last 700 ka". Quaternary Science Reviews. 223: 105950. Bibcode:2019QSRv..22305950M. doi:10.1016/j.quascirev.2019.105950.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Stuart, A. J. & Lister, A. M. (2011). "Extinction chronology of the cave lion Panthera spelaea". Quaternary Science Reviews. 30 (17): 2329–2340. Bibcode:2011QSRv...30.2329S. doi:10.1016/j.quascirev.2010.04.023.
^ Goldfuss, G. A. (1810). Die Umgebungen von Muggensdorf. Erlangen: Johann Jakob Palm.
^ Diedrich, C. G. (2008). "The holotypes of the upper Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823: Hyaenidae) and Panthera leo spelaea (Goldfuss, 1810: Felidae) of the Zoolithen Cave hyena den (South Germany) and their palaeo-ecological interpretation". Zoological Journal of the Linnean Society. 154 (4): 822–831. doi:10.1111/j.1096-3642.2008.00425.x.
^ Kurtén, B. (1968). Pleistocene Mammals of Europe. London: Weidenfeld and Nicolson.
^ Hemmer, H. (1974). "Untersuchungen zur Stammesgeschichte der Pantherkatzen (Pantherinae) Teil 3. Zur Artgeschichte des Löwen Panthera (Panthera) leo (Linnaeus, 1758)". Veröffentlichungen der Zoologischen Staatssammlung 17: 167–280.
^ Turner, A. (1984). "Dental sex dimorphism in European lions (Panthera leo L.) of the Upper Pleistocene: palaeoecological and palaeoethological implications". Annales Zoologici Fennici. 21: 1–8.
^ Groiss, J. Th. (1996). "Der Höhlentiger Panthera tigris spelaea (Goldfuss)". Neues Jahrbuch für Geologie und Paläontologie (7): 399–414. doi:10.1127/njgpm/1996/1996/399.
^ Spassov, N. & Iliev, N. (1994). "Animal remains from the submerged Late Eneolithic – early Bronze Age settlements in Sozopol (South Bulgarian Black Sea Coast)". Proceedings of the International Symposium VI. Thracia Pontica. pp. 287–314.{{cite book}}: CS1 maint: location missing publisher (link)
^ Sotnikova, M.; Nikolskiy, P. (2006). "Systematic position of the cave lion Panthera spelaea (Goldfuss) based on cranial and dental characters" (PDF). Quaternary International. 142–143: 218–228. Bibcode:2006QuInt.142..218S. doi:10.1016/j.quaint.2005.03.019. Archived (PDF) from the original on 5 February 2023. Retrieved 22 January 2023.
^ Christiansen, P. (2008). "Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters". Cladistics. 24 (6): 977–992. doi:10.1111/j.1096-0031.2008.00226.x. PMID 34892880. S2CID 84497516.
^ ^a ^b ^c Barnett, R.; Shapiro, B.; Barnes, I. A. N.; Ho, S. Y.; Burger, J.; Yamaguchi, N.; Higham, T. F.; Wheeler, H.; Rosendahl, W.; Sher, A. V. & Sotnikova, M. (2009). "Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity" (PDF). Molecular Ecology. 18 (8): 1668–1677. Bibcode:2009MolEc..18.1668B. doi:10.1111/j.1365-294X.2009.04134.x. PMID 19302360. S2CID 46716748. Archived from the original (PDF) on 21 January 2012.
^ ^a ^b Barnett, R.; Mendoza, M. L. Z.; Soares, A. E. R.; Ho, S. Y. W.; Zazula, G.; Yamaguchi, N.; Shapiro, B.; Kirillova, I. V.; Larson, G.; Gilbert, M. T. P. (2016). "Mitogenomics of the Extinct Cave Lion, Panthera spelaea (Goldfuss, 1810), Resolve its Position within the Panthera Cats". Open Quaternary. 2: 4. doi:10.5334/oq.24. hdl:10576/22920.
^ Baryshnikov, G. F. & Boeskorov, G. (2001). "The Pleistocene cave lion, Panthera spelaea (Carnivora, Felidae) from Yakutia, Russia". Cranium. 18 (1): 7–24. Archived from the original on 1 January 2018. Retrieved 31 December 2017.
^ ^a ^b Stanton, D.W.; Alberti, F.; Plotnikov, V.; Androsov, S.; Grigoriev, S.; Fedorov, S.; Kosintsev, P.; Nagel, D.; Vartanyan, S.; Barnes, I. & Barnett, R. (2020). "Early Pleistocene origin and extensive intra-species diversity of the extinct cave lion". Scientific Reports. 10 (1): 12621. Bibcode:2020NatSR..1012621S. doi:10.1038/s41598-020-69474-1. PMC 7387438. PMID 32724178.
^ ^a ^b ^c ^d ^e ^f ^g Boeskorov, G. G.; Plotnikov, V. V.; Protopopov, A. V.; Baryshnikov, G. F.; Fosse, P.; Dalén, L.; David, S. W. G.; Pavlov, I. S.; Naoki, S. & Alexey, N. T. (2021). "The preliminary analysis of Cave Lion cubs Panthera spelaea (Goldfuss, 1810) from the permafrost of Siberia". Quaternary. 4 (3): 24. doi:10.3390/quat4030024.
^ ^a ^b Sotnikova, M.V. & Foronova, I.V. (2014). "First Asian record of Panthera (Leo) fossilis (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia". Integrative Zoology. 9 (4): 517–530. doi:10.1111/1749-4877.12082. PMID 24382145.
^ ^a ^b Marciszak, Adrian; Lipecki, Grzegorz; Pawłowska, Kamilla; Jakubowski, Gwidon; Ratajczak-Skrzatek, Urszula; Zarzecka-Szubińska, Katarzyna; Nadachowski, Adam (20 December 2021). "The Pleistocene lion Panthera spelaea (Goldfuss, 1810) from Poland – A review". Quaternary International. The Quaternary of Europe and Adjacent Areas: Stratigraphical Perspectives and Tools for Correlations – SEQS-2019. 605–606: 213–240. Bibcode:2021QuInt.605..213M. doi:10.1016/j.quaint.2020.12.018. ISSN 1040-6182. Archived from the original on 18 May 2024. Retrieved 22 March 2024 – via Elsevier Science Direct.
^ ^a ^b Iannucci, A.; Mecozzi, B.; Pineda, A.; Raffaele, S.; Carpentieri, M.; Rabinovich, R.; Moncel, M.-H. (24 June 2024). "Early occurrence of lion (Panthera spelaea) at the Middle Pleistocene Acheulean site of Notarchirico (MIS 16, Italy)". Quaternary Science. 39 (5): 683–690. Bibcode:2024JQS....39..683I. doi:10.1002/jqs.3639. ISSN 0267-8179.
^ Sotnikova, Marina V.; Foronova, Irina V. (August 2014). "First Asian record of Panthera (Leo) fossilis (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia". Integrative Zoology. 9 (4): 517–530. doi:10.1111/1749-4877.12082. ISSN 1749-4877.
^ Sabo, Martin; Tomašových, Adam; Gullár, Juraj (August 2022). "Geographic and temporal variability in Pleistocene lion-like felids: Implications for their evolution and taxonomy". Palaeontologia Electronica. 25 (2): 1–27. doi:10.26879/1175. ISSN 1094-8074. S2CID 251855356.
^ ^a ^b Marciszak, Adrian; Ivanoff, Dmitry V.; Semenov, Yuriy A.; Talamo, Sahra; Ridush, Bogdan; Stupak, Alina; Yanish, Yevheniia; Kovalchuk, Oleksandr (March 2023). "The Quaternary lions of Ukraine and a trend of decreasing size in Panthera spelaea". Journal of Mammalian Evolution. 30 (1): 109–135. doi:10.1007/s10914-022-09635-3. ISSN 1064-7554.
^ Persico, Davide (June 2021). "First fossil record of cave lion (Panthera (Leo) spelaea intermedia) from alluvial deposits of the Po River in northern Italy". Quaternary International. 586: 14–23. Bibcode:2021QuInt.586...14P. doi:10.1016/j.quaint.2021.02.029.
^ Salis, Alexander T.; Bray, Sarah C. E.; Lee, Michael S. Y.; Heiniger, Holly; Barnett, Ross; Burns, James A.; Doronichev, Vladimir; Fedje, Daryl; Golovanova, Liubov; Harington, C. Richard; Hockett, Bryan; Kosintsev, Pavel; Lai, Xulong; Mackie, Quentin; Vasiliev, Sergei (December 2022). "Lions and brown bears colonized North America in multiple synchronous waves of dispersal across the Bering Land Bridge". Molecular Ecology. 31 (24): 6407–6421. Bibcode:2022MolEc..31.6407S. doi:10.1111/mec.16267. hdl:11343/299180. ISSN 0962-1083. PMID 34748674.
^ ^a ^b Chauvet, J.-M.; Brunel, D. E. & Hillaire, C. (1996). Dawn of Art: The Chauvet Cave. The oldest known paintings in the world. New York: Harry N. Abrams.
^ Leroi-Gourhan, A.; Allain, J. (1979). Lascaux inconnu. XXIIe supplement à "Gallia Préhistoire". Paris.{{cite book}}: CS1 maint: location missing publisher (link)
^ Yamaguchi, N.; Cooper, A.; Werdelin, L.; MacDonald, D. W. (2004). "Evolution of the mane and group-living in the lion (Panthera leo): a review". Journal of Zoology. 263 (4): 329–342. doi:10.1017/S0952836904005242.
^ Marciszak, Adrian; Gornig, Wiktoria (September 2024). "From giant to dwarf: A trend of decreasing size in Panthera spelaea (Goldfuss, 1810) and its likely implications". Earth History and Biodiversity. 1: 100007. doi:10.1016/j.hisbio.2024.100007.
^ Diedrich, C. G. (2011). "Late Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeletons from the Czech Republic (central Europe); their pathological cranial features and injuries resulting from intraspecific fights, conflicts with hyenas, and attacks on cave bears". Bulletin of Geosciences. 86 (4): 817–840. doi:10.3140/bull.geosci.1263.
^ Chernova, O. F.; Kirillova, I. V.; Shapiro, B.; Shidlovskiy, F. K.; Soares, A. E. R.; Levchenko, V. A.; Bertuch, F. (2016). "Morphological and genetic identification and isotopic study of the hair of a cave lion (Panthera spelaea Goldfuss, 1810) from the Malyi Anyui River (Chukotka, Russia)". Quaternary Science Reviews. 142: 61–73. Bibcode:2016QSRv..142...61C. doi:10.1016/j.quascirev.2016.04.018.
^ Ersmark, E.; Orlando, L.; Sandoval-Castellanos, E.; Barnes, I.; Barnett, R.; Stuart, A.; Lister, A.; Dalén, L. (2015). "Population Demography and Genetic Diversity in the Pleistocene Cave Lion". Open Quaternary. 1 (1): Art. 4. doi:10.5334/oq.aa.
^ Álvarez-Lao, D. J.; Álvarez-Vena, A.; Ballesteros, D.; García, N.; Laplana, C. (2020). "A cave lion (Panthera spelaea) skeleton from Torca del León (NW Iberia): Micromammals indicate a temperate and forest environment corresponding to GI-11 (MIS 3)". Quaternary Science Reviews. 229: 106123. Bibcode:2020QSRv..22906123A. doi:10.1016/j.quascirev.2019.106123. hdl:10651/54312.
^ Iannucci, Alessio, Mecozzi, Beniamino, Pineda, Antonio, Sardella, Raffaele, Carpentieri, Marco, Rabinovich, Rivka, and Moncel Marie-Helene. "Early Occurrence of lion (Panthera spelaea) at the Middle Pleistocene Acheulean site of Notarchirico (MIS 16, Italy). Journal of Quarternary Sciences 39:3 (2024): 4.
^ Smith, Geoff M.; Spasov, Rosen; Martisius, Naomi L.; Sinet-Mathiot, Virginie; Aldeias, Vera; Rezek, Zeljko; Ruebens, Karen; Pederzani, Sarah; McPherron, Shannon P.; Sirakova, Svoboda; Sirakov, Nikolay; Tsanova, Tsenka; Hublin, Jean-Jacques (December 2021). "Subsistence behavior during the Initial Upper Paleolithic in Europe: Site use, dietary practice, and carnivore exploitation at Bacho Kiro Cave (Bulgaria)". Journal of Human Evolution. 161: 103074. Bibcode:2021JHumE.16103074S. doi:10.1016/j.jhevol.2021.103074. PMID 34628301. Retrieved 22 March 2024 – via Elsevier Science Direct.
^ Sabol, Martin; Döppes, Doris (January 2020). "The first radiocarbon dating of a cave lion fossil from the Slovakian Western Carpathians". Acta Geologica Slovaca. 11 (1): 11–14 – via ResearchGate.
^ Diedrich, Cajus G.; Rathgeber, Thomas (12 October 2011). "Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss 1810) skeleton remains of the Upper Rhine Valley (SW Germany) and contributions to their sexual dimorphism, taphonomy and habitus". Historical Biology. 24 (1): 1–28. doi:10.1080/08912963.2010.549943. ISSN 0891-2963. Archived from the original on 18 May 2024. Retrieved 24 March 2024 – via Taylor and Francis Online.
^ Marciszak, Adrian; Schouwenburg, Charles; Gornig, Wiktoria; Lipecki, Grzegorz; Mackiewicz, Paweł (1 November 2019). "Morphometric comparison of Panthera spelaea (Goldfuss, 1810) from Poland with the lion remains from Eurasia over the last 700 ka". Quaternary Science Reviews. 223: 105950. Bibcode:2019QSRv..22305950M. doi:10.1016/j.quascirev.2019.105950. Archived from the original on 29 March 2023. Retrieved 24 March 2024 – via Elsevier Science Direct.
^ Gimranov, Dmitry; Kosintsev, Pavel (30 April 2020). "Quaternary large mammals from the Imanay Cave". Quaternary International. Quaternary Stratigraphy and Karst & Cave Sediments: the INQUA-SEQS 2018 Meeting. 546: 125–134. Bibcode:2020QuInt.546..125G. doi:10.1016/j.quaint.2020.01.014. ISSN 1040-6182. Archived from the original on 18 May 2024. Retrieved 22 March 2024 – via Elsevier Science Direct.
^ Sherani, Shaheer; Perng, Liongvi; Sherani, Maryam (3 June 2023). "Evidence of cave lion (Panthera spelaea) from Pleistocene Northeast China". Historical Biology. 35 (6): 988–996. Bibcode:2023HBio...35..988S. doi:10.1080/08912963.2022.2071711. ISSN 0891-2963.
^ Puzachenko, A. Yu.; Argant, A.; Baryshnikov, G. F.; Boeskorov, G. G.; Kirillova, I. V.; Klimovsky, A. I.; Kosintsev, P. A.; Marciszak, A.; Plotnikov, V. V.; Sabol, M.; Vasiliev, S. K. (2024). "Distribution history of the cave lion (Panthera spelaea (Goldfuss, 1810))". Earth History and Biodiversity. 1. 100006. doi:10.1016/j.hisbio.2024.100006.
^ Sabol, Martin; Puzachenko, Andrey Yu. (September 2024). "Distribution of cave lions (Panthera spelaea ssp.) in mountain areas". Earth History and Biodiversity. 1: 100009. doi:10.1016/j.hisbio.2024.100009.
^ Sabol, M.; Gullár, J.; Horvát, J. (2018). "Montane record of the late Pleistocene Panthera spelaea (Goldfuss, 1810) from the Západné Tatry Mountains (northern Slovakia)". Journal of Vertebrate Paleontology. 38 (3): e1467921. Bibcode:2018JVPal..38E7921S. doi:10.1080/02724634.2018.1467921.
^ Álvarez-Lao, Diego J.; Álvarez-Vena, Adrián; Ballesteros, Daniel; García, Nuria; Laplana, César (February 2020). "A cave lion (Panthera spelaea) skeleton from Torca del León (NW Iberia): Micromammals indicate a temperate and forest environment corresponding to GI-11 (MIS 3)". Quaternary Science Reviews. 229: 106123. Bibcode:2020QSRv..22906123A. doi:10.1016/j.quascirev.2019.106123. hdl:10651/54312.
^ Diedrich, Cajus G. (June 2011). "A diseased Panthera leo spelaea (Goldfuss 1810) lioness from a forest elephant graveyard in the Late Pleistocene (Eemian) interglacial lake at Neumark-Nord, central Germany". Historical Biology. 23 (2–3): 195–217. Bibcode:2011HBio...23..195D. doi:10.1080/08912963.2010.507814. ISSN 0891-2963.
^ Bölsche, W. & Harder, H. (1900). Tiere der Urwelt. Serie III. Wandsbek-Hamburg: Verlag der Kakao-Compagnie Theodor Reichardt.
^ ^a ^b ^c ^d ^e ^f ^g Bocherens, H. (2015). "Isotopic tracking of large carnivore palaeoecology in the mammoth steppe". Quaternary Science Reviews. 117: 42–71. Bibcode:2015QSRv..117...42B. doi:10.1016/j.quascirev.2015.03.018.
^ Diedrich, C. G. (2011). "The largest European lion Panthera leo spelaea (Goldfuss 1810) population from the Zoolithen Cave, Germany: specialised cave bear predators of Europe". Historical Biology. 23 (2–3): 271–311. Bibcode:2011HBio...23..271D. doi:10.1080/08912963.2010.546529. S2CID 86638786.
^ Diedrich, C. G. (2011). "Pleistocene Panthera leo spelaea (Goldfuss 1810) remains from the Balve cave (NW Germany) - a cave bear, hyena den and middle palaeolithic human cave – and review of the Sauerland Karst lion cave sites". Quaternaire. 22 (2): 105–127. doi:10.4000/quaternaire.5897. Archived from the original on 11 April 2021. Retrieved 18 September 2020.
^ ^a ^b Diedrich, C.G. & ŽÁK, K. 2006. Prey deposits and den sites of the Upper Pleistocene hyena Crocuta crocuta spelaea (Goldfuss, 1823) in horizontal and vertical caves of the Bohemian Karst (Czech Republic). Bulletin of Geosciences 81(4), 237–276 (25 figures). Czech Geological Survey, Prague. ISSN 1214-1119.
^ Diedrich, Cajus G. (1 July 2011). "Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss, 1810) footprints and bone records from open air sites in northern Germany – Evidence of hyena-lion antagonism and scavenging in Europe". Quaternary Science Reviews. 30 (15): 1883–1906. Bibcode:2011QSRv...30.1883D. doi:10.1016/j.quascirev.2011.03.006. ISSN 0277-3791. Archived from the original on 18 May 2024. Retrieved 4 February 2024 – via Elsevier Science Direct.
^ ^a ^b Bocherens, H.; Drucker, D. G.; Bonjean, D.; Bridault, A.; Conard, N. J.; Cupillard, C.; Germonpré, M.; Höneisen, M.; Münzel, S. C.; Napierala, H. & Patou-Mathis, M. (2011). "Isotopic evidence for dietary ecology of cave lion (Panthera spelaea) in North-Western Europe: prey choice, competition and implications for extinction" (PDF). Quaternary International. 245 (2): 249–261. Bibcode:2011QuInt.245..249B. doi:10.1016/j.quaint.2011.02.023. S2CID 129706533. Archived (PDF) from the original on 11 May 2020. Retrieved 18 September 2020.
^ Drucker, Dorothée G. (31 May 2022). "The Isotopic Ecology of the Mammoth Steppe". Annual Review of Earth and Planetary Sciences. 50 (1): 395–418. doi:10.1146/annurev-earth-100821-081832. ISSN 0084-6597.
^ Diedrich, C. G. (2013). "Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art". Quaternary Science Reviews. 76: 167–193. Bibcode:2013QSRv...76..167D. doi:10.1016/j.quascirev.2013.05.009.
^ ^a ^b Russo, G.; Milks, A.; Leder, D.; Koddenberg, T.; Starkovich, B. M.; Duval, M.; Zhao, J.-X.; Darga, R.; Rosendahl, W.; Terberger, T. (2023). "First direct evidence of lion hunting and the early use of a lion pelt by Neanderthals". Scientific Reports. 13 (1): 16405. Bibcode:2023NatSR..1316405R. doi:10.1038/s41598-023-42764-0. PMC 10570355. PMID 37828055.
^ Cueto, M.; Camarós, E.; Castaños, P.; Ontañón, R.; Arias, P. (2016). "Under the skin of a Lion: Unique evidence of Upper Paleolithic exploitation and use of Cave Lion (Panthera spelaea) from the Lower Gallery of La Garma (Spain)". PLOS ONE. 11 (10): e0163591. Bibcode:2016PLoSO..1163591C. doi:10.1371/journal.pone.0163591. PMC 5082676. PMID 27783697.
^ Sigari, D.; Bourdier, C.; Conti, C.; Conti, J.; Forti, L.; García-Diez, M.; Lai, G.; Mazzini, I.; Pieruccini, P.; Sardella, R. (15 June 2024). "The last cave lion of the late Upper Palaeolithic: The engraved feline of Grotta Romanelli (southern Italy)". Quaternary Science Reviews. 334: 108670. doi:10.1016/j.quascirev.2024.108670.
^ Ersmark, Erik; Orlando, Ludovic; Sandoval-Castellanos, Edson; Barnes, Ian; Barnett, Ross; Stuart, Anthony; Lister, Adrian; Dalén, Love (9 March 2015). "Population Demography and Genetic Diversity in the Pleistocene Cave Lion". Open Quaternary. 1 (1): 4. doi:10.5334/oq.aa. ISSN 2055-298X.
^ Kirillova, I. V.; Tiunov, A. V.; Levchenko, V. A.; Chernova, O. F.; Yudin, V. G.; Bertuch, F.; Shidlovskiy, F. K. (2015). "On the discovery of a cave lion from the Malyi Anyui River (Chukotka, Russia)". Quaternary Science Reviews. 117: 135–151. Bibcode:2015QSRv..117..135K. doi:10.1016/j.quascirev.2015.03.029.
^ "Extinct lion cubs found in Siberia are up to 55,000 years old – latest test results reveal". The Siberian Times. 3 November 2016. Archived from the original on 12 November 2017. Retrieved 12 November 2017.
^ Koryakina, A. (2015). "Meet this extinct cave lion, at least 10,000 years old – world exclusive". The Siberian Times. Archived from the original on 17 April 2019. Retrieved 29 October 2015.
^ Black, R. (2015). "Frozen Cave Lion Cubs from the Ice Age Found in Siberia". National Geographic News. National Geographic. Archived from the original on 26 February 2021. Retrieved 29 October 2015.
^ "Whiskers still bristling after more than 12,000 years in the Siberian cold". The Siberian Times. 2015. Archived from the original on 16 December 2016. Retrieved 25 November 2016.
^ "Extinct cave lion cub in 'perfect' condition found in Siberia rising cloning hopes". The Siberian Times. 2017. Archived from the original on 12 November 2017. Retrieved 12 November 2017.
^ Gertcyk, O. (2018). "Cute first pictures of new 50,000 year old cave lion cub found perfectly preserved in permafrost". The Siberian Times. Archived from the original on 14 September 2018. Retrieved 16 September 2018.
^ Gertcyk, O. (9 August 2021). "Meet Sparta, the 'best preserved ice age animal ever found'". The Siberian Times. Archived from the original on 12 August 2021. Retrieved 13 August 2021.

External links

Prehistoric cats and prehistoric cat-like creatures, from the Messybeast Cat Resource Archive
The mammoth and the flood, volume 5, chapter 1, by Hans Krause.
Hoyle and cavetigers Archived 13 January 2016 at the Wayback Machine, from the Dinosaur Mailing List. (Groiss)
Photo-reconstruction of Panthera spelaea, by paleoartist Roman Uchytel

[Burger-1] Burger, J.; Rosendahl, W.; Loreille, O.; Hemmer, H.; Eriksson, T.; Götherström, A.; Hiller, J.; Collins, M. J.; Wess, T.; Alt, K. W. (2004). "Molecular phylogeny of the extinct cave lion Panthera leo spelaea". Molecular Phylogenetics and Evolution. 30 (3): 841–849. Bibcode:2004MolPE..30..841B. doi:10.1016/j.ympev.2003.07.020. PMID 15012963.

[Manuel_al2020-2] Manuel, M. d.; Ross, B.; Sandoval-Velasco, M.; Yamaguchi, N.; Vieira, F. G.; Mendoza, M. L. Z.; Liu, S.; Martin, M. D.; Sinding, M.-H. S.; Mak, S. S. T.; Carøe, C.; Liu, S.; Guo, C.; Zheng, J.; Zazula, G.; Baryshnikov, G.; Eizirik, E.; Koepfli, K.-P.; Johnson, W. E.; Antunes, A.; Sicheritz-Ponten, T.; Gopalakrishnan, S.; Larson, G.; Yang, H.; O'Brien, S. J.; Hansen, A. J.; Zhang, G.; Marques-Bonet, T. & Gilbert, M. T. P. (2020). "The evolutionary history of extinct and living lions". Proceedings of the National Academy of Sciences of the United States of America. 117 (20): 10927–10934. Bibcode:2020PNAS..11710927D. doi:10.1073/pnas.1919423117. PMC 7245068. PMID 32366643.

[:2-3] Marciszak, Adrian; Schouwenburg, Charles; Gornig, Wiktoria; Lipecki, Grzegorz; Mackiewicz, Paweł (November 2019). "Morphometric comparison of Panthera spelaea (Goldfuss, 1810) from Poland with the lion remains from Eurasia over the last 700 ka". Quaternary Science Reviews. 223: 105950. Bibcode:2019QSRv..22305950M. doi:10.1016/j.quascirev.2019.105950.

[Stuart2011-4] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Stuart, A. J. & Lister, A. M. (2011). "Extinction chronology of the cave lion Panthera spelaea". Quaternary Science Reviews. 30 (17): 2329–2340. Bibcode:2011QSRv...30.2329S. doi:10.1016/j.quascirev.2010.04.023.

[5] Goldfuss, G. A. (1810). Die Umgebungen von Muggensdorf. Erlangen: Johann Jakob Palm.

[6] Diedrich, C. G. (2008). "The holotypes of the upper Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823: Hyaenidae) and Panthera leo spelaea (Goldfuss, 1810: Felidae) of the Zoolithen Cave hyena den (South Germany) and their palaeo-ecological interpretation". Zoological Journal of the Linnean Society. 154 (4): 822–831. doi:10.1111/j.1096-3642.2008.00425.x.

[7] Kurtén, B. (1968). Pleistocene Mammals of Europe. London: Weidenfeld and Nicolson.

[Hemmer-8] Hemmer, H. (1974). "Untersuchungen zur Stammesgeschichte der Pantherkatzen (Pantherinae) Teil 3. Zur Artgeschichte des Löwen Panthera (Panthera) leo (Linnaeus, 1758)". Veröffentlichungen der Zoologischen Staatssammlung 17: 167–280.

[9] Turner, A. (1984). "Dental sex dimorphism in European lions (Panthera leo L.) of the Upper Pleistocene: palaeoecological and palaeoethological implications". Annales Zoologici Fennici. 21: 1–8.

[10] Groiss, J. Th. (1996). "Der Höhlentiger Panthera tigris spelaea (Goldfuss)". Neues Jahrbuch für Geologie und Paläontologie (7): 399–414. doi:10.1127/njgpm/1996/1996/399.

[11] Spassov, N. & Iliev, N. (1994). "Animal remains from the submerged Late Eneolithic – early Bronze Age settlements in Sozopol (South Bulgarian Black Sea Coast)". Proceedings of the International Symposium VI. Thracia Pontica. pp. 287–314.{{cite book}}: CS1 maint: location missing publisher (link)

[12] Sotnikova, M.; Nikolskiy, P. (2006). "Systematic position of the cave lion Panthera spelaea (Goldfuss) based on cranial and dental characters" (PDF). Quaternary International. 142–143: 218–228. Bibcode:2006QuInt.142..218S. doi:10.1016/j.quaint.2005.03.019. Archived (PDF) from the original on 5 February 2023. Retrieved 22 January 2023.

[13] Christiansen, P. (2008). "Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters". Cladistics. 24 (6): 977–992. doi:10.1111/j.1096-0031.2008.00226.x. PMID 34892880. S2CID 84497516.

[Barnett2009-14] Barnett, R.; Shapiro, B.; Barnes, I. A. N.; Ho, S. Y.; Burger, J.; Yamaguchi, N.; Higham, T. F.; Wheeler, H.; Rosendahl, W.; Sher, A. V. & Sotnikova, M. (2009). "Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity" (PDF). Molecular Ecology. 18 (8): 1668–1677. Bibcode:2009MolEc..18.1668B. doi:10.1111/j.1365-294X.2009.04134.x. PMID 19302360. S2CID 46716748. Archived from the original (PDF) on 21 January 2012.

[Barnett2016-15] Barnett, R.; Mendoza, M. L. Z.; Soares, A. E. R.; Ho, S. Y. W.; Zazula, G.; Yamaguchi, N.; Shapiro, B.; Kirillova, I. V.; Larson, G.; Gilbert, M. T. P. (2016). "Mitogenomics of the Extinct Cave Lion, Panthera spelaea (Goldfuss, 1810), Resolve its Position within the Panthera Cats". Open Quaternary. 2: 4. doi:10.5334/oq.24. hdl:10576/22920.

[16] Baryshnikov, G. F. & Boeskorov, G. (2001). "The Pleistocene cave lion, Panthera spelaea (Carnivora, Felidae) from Yakutia, Russia". Cranium. 18 (1): 7–24. Archived from the original on 1 January 2018. Retrieved 31 December 2017.

[Stanton2020-17] Stanton, D.W.; Alberti, F.; Plotnikov, V.; Androsov, S.; Grigoriev, S.; Fedorov, S.; Kosintsev, P.; Nagel, D.; Vartanyan, S.; Barnes, I. & Barnett, R. (2020). "Early Pleistocene origin and extensive intra-species diversity of the extinct cave lion". Scientific Reports. 10 (1): 12621. Bibcode:2020NatSR..1012621S. doi:10.1038/s41598-020-69474-1. PMC 7387438. PMID 32724178.

[Boeskorov_al.2021-18] ^ ^a ^b ^c ^d ^e ^f ^g Boeskorov, G. G.; Plotnikov, V. V.; Protopopov, A. V.; Baryshnikov, G. F.; Fosse, P.; Dalén, L.; David, S. W. G.; Pavlov, I. S.; Naoki, S. & Alexey, N. T. (2021). "The preliminary analysis of Cave Lion cubs Panthera spelaea (Goldfuss, 1810) from the permafrost of Siberia". Quaternary. 4 (3): 24. doi:10.3390/quat4030024.

[Sotnikova2014-19] Sotnikova, M.V. & Foronova, I.V. (2014). "First Asian record of Panthera (Leo) fossilis (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia". Integrative Zoology. 9 (4): 517–530. doi:10.1111/1749-4877.12082. PMID 24382145.

[:0-20] Marciszak, Adrian; Lipecki, Grzegorz; Pawłowska, Kamilla; Jakubowski, Gwidon; Ratajczak-Skrzatek, Urszula; Zarzecka-Szubińska, Katarzyna; Nadachowski, Adam (20 December 2021). "The Pleistocene lion Panthera spelaea (Goldfuss, 1810) from Poland – A review". Quaternary International. The Quaternary of Europe and Adjacent Areas: Stratigraphical Perspectives and Tools for Correlations – SEQS-2019. 605–606: 213–240. Bibcode:2021QuInt.605..213M. doi:10.1016/j.quaint.2020.12.018. ISSN 1040-6182. Archived from the original on 18 May 2024. Retrieved 22 March 2024 – via Elsevier Science Direct.

[Ianucci_al.2024-21] Iannucci, A.; Mecozzi, B.; Pineda, A.; Raffaele, S.; Carpentieri, M.; Rabinovich, R.; Moncel, M.-H. (24 June 2024). "Early occurrence of lion (Panthera spelaea) at the Middle Pleistocene Acheulean site of Notarchirico (MIS 16, Italy)". Quaternary Science. 39 (5): 683–690. Bibcode:2024JQS....39..683I. doi:10.1002/jqs.3639. ISSN 0267-8179.

[22] Sotnikova, Marina V.; Foronova, Irina V. (August 2014). "First Asian record of Panthera (Leo) fossilis (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia". Integrative Zoology. 9 (4): 517–530. doi:10.1111/1749-4877.12082. ISSN 1749-4877.

[:02-23] Sabo, Martin; Tomašových, Adam; Gullár, Juraj (August 2022). "Geographic and temporal variability in Pleistocene lion-like felids: Implications for their evolution and taxonomy". Palaeontologia Electronica. 25 (2): 1–27. doi:10.26879/1175. ISSN 1094-8074. S2CID 251855356.

[:1-24] Marciszak, Adrian; Ivanoff, Dmitry V.; Semenov, Yuriy A.; Talamo, Sahra; Ridush, Bogdan; Stupak, Alina; Yanish, Yevheniia; Kovalchuk, Oleksandr (March 2023). "The Quaternary lions of Ukraine and a trend of decreasing size in Panthera spelaea". Journal of Mammalian Evolution. 30 (1): 109–135. doi:10.1007/s10914-022-09635-3. ISSN 1064-7554.

[25] Persico, Davide (June 2021). "First fossil record of cave lion (Panthera (Leo) spelaea intermedia) from alluvial deposits of the Po River in northern Italy". Quaternary International. 586: 14–23. Bibcode:2021QuInt.586...14P. doi:10.1016/j.quaint.2021.02.029.

[26] Salis, Alexander T.; Bray, Sarah C. E.; Lee, Michael S. Y.; Heiniger, Holly; Barnett, Ross; Burns, James A.; Doronichev, Vladimir; Fedje, Daryl; Golovanova, Liubov; Harington, C. Richard; Hockett, Bryan; Kosintsev, Pavel; Lai, Xulong; Mackie, Quentin; Vasiliev, Sergei (December 2022). "Lions and brown bears colonized North America in multiple synchronous waves of dispersal across the Bering Land Bridge". Molecular Ecology. 31 (24): 6407–6421. Bibcode:2022MolEc..31.6407S. doi:10.1111/mec.16267. hdl:11343/299180. ISSN 0962-1083. PMID 34748674.

[Chauvet1996-27] Chauvet, J.-M.; Brunel, D. E. & Hillaire, C. (1996). Dawn of Art: The Chauvet Cave. The oldest known paintings in the world. New York: Harry N. Abrams.

[28] Leroi-Gourhan, A.; Allain, J. (1979). Lascaux inconnu. XXIIe supplement à "Gallia Préhistoire". Paris.{{cite book}}: CS1 maint: location missing publisher (link)

[Yamaguchietal2004-29] Yamaguchi, N.; Cooper, A.; Werdelin, L.; MacDonald, D. W. (2004). "Evolution of the mane and group-living in the lion (Panthera leo): a review". Journal of Zoology. 263 (4): 329–342. doi:10.1017/S0952836904005242.

[30] Marciszak, Adrian; Gornig, Wiktoria (September 2024). "From giant to dwarf: A trend of decreasing size in Panthera spelaea (Goldfuss, 1810) and its likely implications". Earth History and Biodiversity. 1: 100007. doi:10.1016/j.hisbio.2024.100007.

[31] Diedrich, C. G. (2011). "Late Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeletons from the Czech Republic (central Europe); their pathological cranial features and injuries resulting from intraspecific fights, conflicts with hyenas, and attacks on cave bears". Bulletin of Geosciences. 86 (4): 817–840. doi:10.3140/bull.geosci.1263.

[32] Chernova, O. F.; Kirillova, I. V.; Shapiro, B.; Shidlovskiy, F. K.; Soares, A. E. R.; Levchenko, V. A.; Bertuch, F. (2016). "Morphological and genetic identification and isotopic study of the hair of a cave lion (Panthera spelaea Goldfuss, 1810) from the Malyi Anyui River (Chukotka, Russia)". Quaternary Science Reviews. 142: 61–73. Bibcode:2016QSRv..142...61C. doi:10.1016/j.quascirev.2016.04.018.

[33] Ersmark, E.; Orlando, L.; Sandoval-Castellanos, E.; Barnes, I.; Barnett, R.; Stuart, A.; Lister, A.; Dalén, L. (2015). "Population Demography and Genetic Diversity in the Pleistocene Cave Lion". Open Quaternary. 1 (1): Art. 4. doi:10.5334/oq.aa.

[34] Álvarez-Lao, D. J.; Álvarez-Vena, A.; Ballesteros, D.; García, N.; Laplana, C. (2020). "A cave lion (Panthera spelaea) skeleton from Torca del León (NW Iberia): Micromammals indicate a temperate and forest environment corresponding to GI-11 (MIS 3)". Quaternary Science Reviews. 229: 106123. Bibcode:2020QSRv..22906123A. doi:10.1016/j.quascirev.2019.106123. hdl:10651/54312.

[35] Iannucci, Alessio, Mecozzi, Beniamino, Pineda, Antonio, Sardella, Raffaele, Carpentieri, Marco, Rabinovich, Rivka, and Moncel Marie-Helene. "Early Occurrence of lion (Panthera spelaea) at the Middle Pleistocene Acheulean site of Notarchirico (MIS 16, Italy). Journal of Quarternary Sciences 39:3 (2024): 4.

[36] Smith, Geoff M.; Spasov, Rosen; Martisius, Naomi L.; Sinet-Mathiot, Virginie; Aldeias, Vera; Rezek, Zeljko; Ruebens, Karen; Pederzani, Sarah; McPherron, Shannon P.; Sirakova, Svoboda; Sirakov, Nikolay; Tsanova, Tsenka; Hublin, Jean-Jacques (December 2021). "Subsistence behavior during the Initial Upper Paleolithic in Europe: Site use, dietary practice, and carnivore exploitation at Bacho Kiro Cave (Bulgaria)". Journal of Human Evolution. 161: 103074. Bibcode:2021JHumE.16103074S. doi:10.1016/j.jhevol.2021.103074. PMID 34628301. Retrieved 22 March 2024 – via Elsevier Science Direct.

[37] Sabol, Martin; Döppes, Doris (January 2020). "The first radiocarbon dating of a cave lion fossil from the Slovakian Western Carpathians". Acta Geologica Slovaca. 11 (1): 11–14 – via ResearchGate.

[38] Diedrich, Cajus G.; Rathgeber, Thomas (12 October 2011). "Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss 1810) skeleton remains of the Upper Rhine Valley (SW Germany) and contributions to their sexual dimorphism, taphonomy and habitus". Historical Biology. 24 (1): 1–28. doi:10.1080/08912963.2010.549943. ISSN 0891-2963. Archived from the original on 18 May 2024. Retrieved 24 March 2024 – via Taylor and Francis Online.

[39] Marciszak, Adrian; Schouwenburg, Charles; Gornig, Wiktoria; Lipecki, Grzegorz; Mackiewicz, Paweł (1 November 2019). "Morphometric comparison of Panthera spelaea (Goldfuss, 1810) from Poland with the lion remains from Eurasia over the last 700 ka". Quaternary Science Reviews. 223: 105950. Bibcode:2019QSRv..22305950M. doi:10.1016/j.quascirev.2019.105950. Archived from the original on 29 March 2023. Retrieved 24 March 2024 – via Elsevier Science Direct.

[40] Gimranov, Dmitry; Kosintsev, Pavel (30 April 2020). "Quaternary large mammals from the Imanay Cave". Quaternary International. Quaternary Stratigraphy and Karst & Cave Sediments: the INQUA-SEQS 2018 Meeting. 546: 125–134. Bibcode:2020QuInt.546..125G. doi:10.1016/j.quaint.2020.01.014. ISSN 1040-6182. Archived from the original on 18 May 2024. Retrieved 22 March 2024 – via Elsevier Science Direct.

[41] Sherani, Shaheer; Perng, Liongvi; Sherani, Maryam (3 June 2023). "Evidence of cave lion (Panthera spelaea) from Pleistocene Northeast China". Historical Biology. 35 (6): 988–996. Bibcode:2023HBio...35..988S. doi:10.1080/08912963.2022.2071711. ISSN 0891-2963.

[42] Puzachenko, A. Yu.; Argant, A.; Baryshnikov, G. F.; Boeskorov, G. G.; Kirillova, I. V.; Klimovsky, A. I.; Kosintsev, P. A.; Marciszak, A.; Plotnikov, V. V.; Sabol, M.; Vasiliev, S. K. (2024). "Distribution history of the cave lion (Panthera spelaea (Goldfuss, 1810))". Earth History and Biodiversity. 1. 100006. doi:10.1016/j.hisbio.2024.100006.

[43] Sabol, Martin; Puzachenko, Andrey Yu. (September 2024). "Distribution of cave lions (Panthera spelaea ssp.) in mountain areas". Earth History and Biodiversity. 1: 100009. doi:10.1016/j.hisbio.2024.100009.

[44] Sabol, M.; Gullár, J.; Horvát, J. (2018). "Montane record of the late Pleistocene Panthera spelaea (Goldfuss, 1810) from the Západné Tatry Mountains (northern Slovakia)". Journal of Vertebrate Paleontology. 38 (3): e1467921. Bibcode:2018JVPal..38E7921S. doi:10.1080/02724634.2018.1467921.

[45] Álvarez-Lao, Diego J.; Álvarez-Vena, Adrián; Ballesteros, Daniel; García, Nuria; Laplana, César (February 2020). "A cave lion (Panthera spelaea) skeleton from Torca del León (NW Iberia): Micromammals indicate a temperate and forest environment corresponding to GI-11 (MIS 3)". Quaternary Science Reviews. 229: 106123. Bibcode:2020QSRv..22906123A. doi:10.1016/j.quascirev.2019.106123. hdl:10651/54312.

[46] Diedrich, Cajus G. (June 2011). "A diseased Panthera leo spelaea (Goldfuss 1810) lioness from a forest elephant graveyard in the Late Pleistocene (Eemian) interglacial lake at Neumark-Nord, central Germany". Historical Biology. 23 (2–3): 195–217. Bibcode:2011HBio...23..195D. doi:10.1080/08912963.2010.507814. ISSN 0891-2963.

[47] Bölsche, W. & Harder, H. (1900). Tiere der Urwelt. Serie III. Wandsbek-Hamburg: Verlag der Kakao-Compagnie Theodor Reichardt.

[Bocherens2015-48] ^ ^a ^b ^c ^d ^e ^f ^g Bocherens, H. (2015). "Isotopic tracking of large carnivore palaeoecology in the mammoth steppe". Quaternary Science Reviews. 117: 42–71. Bibcode:2015QSRv..117...42B. doi:10.1016/j.quascirev.2015.03.018.

[49] Diedrich, C. G. (2011). "The largest European lion Panthera leo spelaea (Goldfuss 1810) population from the Zoolithen Cave, Germany: specialised cave bear predators of Europe". Historical Biology. 23 (2–3): 271–311. Bibcode:2011HBio...23..271D. doi:10.1080/08912963.2010.546529. S2CID 86638786.

[50] Diedrich, C. G. (2011). "Pleistocene Panthera leo spelaea (Goldfuss 1810) remains from the Balve cave (NW Germany) - a cave bear, hyena den and middle palaeolithic human cave – and review of the Sauerland Karst lion cave sites". Quaternaire. 22 (2): 105–127. doi:10.4000/quaternaire.5897. Archived from the original on 11 April 2021. Retrieved 18 September 2020.

[diedrich&zak-51] Diedrich, C.G. & ŽÁK, K. 2006. Prey deposits and den sites of the Upper Pleistocene hyena Crocuta crocuta spelaea (Goldfuss, 1823) in horizontal and vertical caves of the Bohemian Karst (Czech Republic). Bulletin of Geosciences 81(4), 237–276 (25 figures). Czech Geological Survey, Prague. ISSN 1214-1119.

[52] Diedrich, Cajus G. (1 July 2011). "Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss, 1810) footprints and bone records from open air sites in northern Germany – Evidence of hyena-lion antagonism and scavenging in Europe". Quaternary Science Reviews. 30 (15): 1883–1906. Bibcode:2011QSRv...30.1883D. doi:10.1016/j.quascirev.2011.03.006. ISSN 0277-3791. Archived from the original on 18 May 2024. Retrieved 4 February 2024 – via Elsevier Science Direct.

[Bocherens-53] Bocherens, H.; Drucker, D. G.; Bonjean, D.; Bridault, A.; Conard, N. J.; Cupillard, C.; Germonpré, M.; Höneisen, M.; Münzel, S. C.; Napierala, H. & Patou-Mathis, M. (2011). "Isotopic evidence for dietary ecology of cave lion (Panthera spelaea) in North-Western Europe: prey choice, competition and implications for extinction" (PDF). Quaternary International. 245 (2): 249–261. Bibcode:2011QuInt.245..249B. doi:10.1016/j.quaint.2011.02.023. S2CID 129706533. Archived (PDF) from the original on 11 May 2020. Retrieved 18 September 2020.

[54] Drucker, Dorothée G. (31 May 2022). "The Isotopic Ecology of the Mammoth Steppe". Annual Review of Earth and Planetary Sciences. 50 (1): 395–418. doi:10.1146/annurev-earth-100821-081832. ISSN 0084-6597.

[Diedrich_2013-55] Diedrich, C. G. (2013). "Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art". Quaternary Science Reviews. 76: 167–193. Bibcode:2013QSRv...76..167D. doi:10.1016/j.quascirev.2013.05.009.

[Russo-56] Russo, G.; Milks, A.; Leder, D.; Koddenberg, T.; Starkovich, B. M.; Duval, M.; Zhao, J.-X.; Darga, R.; Rosendahl, W.; Terberger, T. (2023). "First direct evidence of lion hunting and the early use of a lion pelt by Neanderthals". Scientific Reports. 13 (1): 16405. Bibcode:2023NatSR..1316405R. doi:10.1038/s41598-023-42764-0. PMC 10570355. PMID 37828055.

[UnderTheSkinOfLion-57] Cueto, M.; Camarós, E.; Castaños, P.; Ontañón, R.; Arias, P. (2016). "Under the skin of a Lion: Unique evidence of Upper Paleolithic exploitation and use of Cave Lion (Panthera spelaea) from the Lower Gallery of La Garma (Spain)". PLOS ONE. 11 (10): e0163591. Bibcode:2016PLoSO..1163591C. doi:10.1371/journal.pone.0163591. PMC 5082676. PMID 27783697.

[58] Sigari, D.; Bourdier, C.; Conti, C.; Conti, J.; Forti, L.; García-Diez, M.; Lai, G.; Mazzini, I.; Pieruccini, P.; Sardella, R. (15 June 2024). "The last cave lion of the late Upper Palaeolithic: The engraved feline of Grotta Romanelli (southern Italy)". Quaternary Science Reviews. 334: 108670. doi:10.1016/j.quascirev.2024.108670.

[59] Ersmark, Erik; Orlando, Ludovic; Sandoval-Castellanos, Edson; Barnes, Ian; Barnett, Ross; Stuart, Anthony; Lister, Adrian; Dalén, Love (9 March 2015). "Population Demography and Genetic Diversity in the Pleistocene Cave Lion". Open Quaternary. 1 (1): 4. doi:10.5334/oq.aa. ISSN 2055-298X.

[60] Kirillova, I. V.; Tiunov, A. V.; Levchenko, V. A.; Chernova, O. F.; Yudin, V. G.; Bertuch, F.; Shidlovskiy, F. K. (2015). "On the discovery of a cave lion from the Malyi Anyui River (Chukotka, Russia)". Quaternary Science Reviews. 117: 135–151. Bibcode:2015QSRv..117..135K. doi:10.1016/j.quascirev.2015.03.029.

[61] "Extinct lion cubs found in Siberia are up to 55,000 years old – latest test results reveal". The Siberian Times. 3 November 2016. Archived from the original on 12 November 2017. Retrieved 12 November 2017.

[62] Koryakina, A. (2015). "Meet this extinct cave lion, at least 10,000 years old – world exclusive". The Siberian Times. Archived from the original on 17 April 2019. Retrieved 29 October 2015.

[63] Black, R. (2015). "Frozen Cave Lion Cubs from the Ice Age Found in Siberia". National Geographic News. National Geographic. Archived from the original on 26 February 2021. Retrieved 29 October 2015.

[64] "Whiskers still bristling after more than 12,000 years in the Siberian cold". The Siberian Times. 2015. Archived from the original on 16 December 2016. Retrieved 25 November 2016.

[65] "Extinct cave lion cub in 'perfect' condition found in Siberia rising cloning hopes". The Siberian Times. 2017. Archived from the original on 12 November 2017. Retrieved 12 November 2017.

[66] Gertcyk, O. (2018). "Cute first pictures of new 50,000 year old cave lion cub found perfectly preserved in permafrost". The Siberian Times. Archived from the original on 14 September 2018. Retrieved 16 September 2018.

[67] Gertcyk, O. (9 August 2021). "Meet Sparta, the 'best preserved ice age animal ever found'". The Siberian Times. Archived from the original on 12 August 2021. Retrieved 13 August 2021.

[1]

[2]

[3]

[4]

[5]

[6]

[7]

[8]

[9]

[10]

[11]

[12]

[13]

[14]

[15]

[16]

[17]

[18]

[19]

[20]

[21]

[22]

[23]

[24]

[25]

[26]

[27]

[28]

[29]

[30]

[31]

[32]

[33]

[34]

[35]

[36]

[37]

[38]

[39]

[40]

[41]

[42]

[43]

[44]

[45]

[46]

[47]

[48]

[49]

[50]

[51]

[52]

[53]

[54]

[55]

[56]

[57]

[58]

[59]

[60]

[61]

[62]

[63]

[64]

[65]

[66]

[67]

@@ Line 17: / Line 17: @@
 }}
-'''''Panthera spelaea''''', commonly known as the '''cave lion''' (or less commonly as the '''steppe lion''') is an [[extinct]] ''[[Panthera]]'' species native to Eurasia and northwest North America during the [[Pleistocene]] epoch. Genetic analysis of [[ancient DNA]] has revealed that while closely related, it was a distinct species genetically isolated from the modern [[lion]] (''Panthera leo''),<ref name="Burger">{{cite journal |last1=Burger |first1=J. |last2=Rosendahl |first2=W. |last3=Loreille |first3=O. |last4=Hemmer |first4=H. |last5=Eriksson |first5=T. |last6=Götherström |first6=A. |last7=Hiller |first7=J. |last8=Collins |first8=M. J. |last9=Wess |first9=T. |last10=Alt |first10=K. W. |title=Molecular phylogeny of the extinct cave lion ''Panthera leo spelaea'' |journal=[[Molecular Phylogenetics and Evolution]] |volume=30 |issue=3 |pages=841–849 |year=2004 |doi=10.1016/j.ympev.2003.07.020 |pmid=15012963|bibcode=2004MolPE..30..841B }}</ref> with the [[genetic divergence]] between the two species estimated at around 500,000 years ago.<ref name="Manuel_al2020">{{cite journal |author1=Manuel, M. d. |author2=Ross, B. |author3=Sandoval-Velasco, M. |author4=Yamaguchi, N. |author5=Vieira, F. G. |author6=Mendoza, M. L. Z. |author7=Liu, S. |author8=Martin, M. D. |author9=Sinding, M.-H. S. |author10=Mak, S. S. T. |author11=Carøe, C. |author12=Liu, S. |author13=Guo, C. |author14=Zheng, J. |author15=Zazula, G. |name-list-style=amp |year=2020 |title=The evolutionary history of extinct and living lions |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=117 |issue=20 |pages=10927–10934 |bibcode=2020PNAS..11710927D |doi=10.1073/pnas.1919423117 |pmc=7245068 |pmid=32366643 |doi-access=free |author16=Baryshnikov, G. |author17=Eizirik, E. |author18=Koepfli, K.-P. |author19=Johnson, W. E. |author20=Antunes, A. |author21=Sicheritz-Ponten, T. |author22=Gopalakrishnan, S. |author23=Larson, G. |author24=Yang, H. |author25=O'Brien, S. J. |author26=Hansen, A. J. |author27=Zhang, G. |author28=Marques-Bonet, T. |author29=Gilbert, M. T. P.}}</ref> The earliest fossils of the ''P. spelaea'' lineage (either regarded as the separate species ''[[Panthera fossilis]]'' or the subspecies ''P. spelaea fossilis'') in Eurasia date to around 700,000 years ago (with possible late Early Pleistocene records).<ref name=":2">{{Cite journal |last1=Marciszak |first1=Adrian |last2=Schouwenburg |first2=Charles |last3=Gornig |first3=Wiktoria |last4=Lipecki |first4=Grzegorz |last5=Mackiewicz |first5=Paweł |date=November 2019 |title=Morphometric comparison of Panthera spelaea (Goldfuss, 1810) from Poland with the lion remains from Eurasia over the last 700 ka |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119303907 |journal=Quaternary Science Reviews |language=en |volume=223 |pages=105950 |doi=10.1016/j.quascirev.2019.105950|bibcode=2019QSRv..22305950M }}</ref> It is closely related and probably ancestral to the [[American lion]] (''Panthera atrox'').<ref name="Manuel_al2020" />  The species ranged from Western Europe to eastern [[Beringia]] in North America, and was a prominent member of the [[mammoth steppe]] fauna, and an important [[apex predator]] across its range. It became extinct about 13,000 years ago.<ref name="Stuart2011">{{cite journal |author1=Stuart, A. J. |author2=Lister, A. M. |name-list-style=amp |year=2011 |title=Extinction chronology of the cave lion ''Panthera spelaea'' |journal=[[Quaternary Science Reviews]] |volume=30 |issue=17 |pages=2329–2340 |bibcode=2011QSRv...30.2329S |doi=10.1016/j.quascirev.2010.04.023}}</ref> It closely resembled living lions with a coat of yellowish-grey fur, though unlike living lions males appear to have lacked manes.
+'''''Panthera spelaea''''', commonly known as the '''cave lion''' (or less commonly as the '''steppe lion'''), is an [[extinct]] ''[[Panthera]]'' species that was native to Eurasia and northwest North America during the [[Pleistocene]] epoch. Genetic analysis of [[ancient DNA]] has revealed that while closely related, it was a distinct species genetically isolated from the modern [[lion]] (''Panthera leo''),<ref name="Burger">{{cite journal |last1=Burger |first1=J. |last2=Rosendahl |first2=W. |last3=Loreille |first3=O. |last4=Hemmer |first4=H. |last5=Eriksson |first5=T. |last6=Götherström |first6=A. |last7=Hiller |first7=J. |last8=Collins |first8=M. J. |last9=Wess |first9=T. |last10=Alt |first10=K. W. |title=Molecular phylogeny of the extinct cave lion ''Panthera leo spelaea'' |journal=[[Molecular Phylogenetics and Evolution]] |volume=30 |issue=3 |pages=841–849 |year=2004 |doi=10.1016/j.ympev.2003.07.020 |pmid=15012963|bibcode=2004MolPE..30..841B }}</ref> with the [[genetic divergence]] between the two species estimated at around 500,000 years ago.<ref name="Manuel_al2020">{{cite journal |author1=Manuel, M. d. |author2=Ross, B. |author3=Sandoval-Velasco, M. |author4=Yamaguchi, N. |author5=Vieira, F. G. |author6=Mendoza, M. L. Z. |author7=Liu, S. |author8=Martin, M. D. |author9=Sinding, M.-H. S. |author10=Mak, S. S. T. |author11=Carøe, C. |author12=Liu, S. |author13=Guo, C. |author14=Zheng, J. |author15=Zazula, G. |name-list-style=amp |year=2020 |title=The evolutionary history of extinct and living lions |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=117 |issue=20 |pages=10927–10934 |bibcode=2020PNAS..11710927D |doi=10.1073/pnas.1919423117 |pmc=7245068 |pmid=32366643 |doi-access=free |author16=Baryshnikov, G. |author17=Eizirik, E. |author18=Koepfli, K.-P. |author19=Johnson, W. E. |author20=Antunes, A. |author21=Sicheritz-Ponten, T. |author22=Gopalakrishnan, S. |author23=Larson, G. |author24=Yang, H. |author25=O'Brien, S. J. |author26=Hansen, A. J. |author27=Zhang, G. |author28=Marques-Bonet, T. |author29=Gilbert, M. T. P.}}</ref> The earliest fossils of the ''P. spelaea'' lineage (either regarded as the separate species ''[[Panthera fossilis]]'' or the subspecies ''P. spelaea fossilis'') in Eurasia date to around 700,000 years ago (with possible late Early Pleistocene records).<ref name=":2">{{Cite journal |last1=Marciszak |first1=Adrian |last2=Schouwenburg |first2=Charles |last3=Gornig |first3=Wiktoria |last4=Lipecki |first4=Grzegorz |last5=Mackiewicz |first5=Paweł |date=November 2019 |title=Morphometric comparison of Panthera spelaea (Goldfuss, 1810) from Poland with the lion remains from Eurasia over the last 700 ka |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119303907 |journal=Quaternary Science Reviews |volume=223 |pages=105950 |doi=10.1016/j.quascirev.2019.105950|bibcode=2019QSRv..22305950M }}</ref> It is closely related and probably ancestral to the [[American lion]] (''Panthera atrox'').<ref name="Manuel_al2020" />  The species ranged from Western Europe to eastern [[Beringia]] in North America, and was a prominent member of the [[mammoth steppe]] fauna, and an important [[apex predator]] across its range. It became extinct about 13,000 years ago.<ref name="Stuart2011">{{cite journal |author1=Stuart, A. J. |author2=Lister, A. M. |name-list-style=amp |year=2011 |title=Extinction chronology of the cave lion ''Panthera spelaea'' |journal=[[Quaternary Science Reviews]] |volume=30 |issue=17 |pages=2329–2340 |bibcode=2011QSRv...30.2329S |doi=10.1016/j.quascirev.2010.04.023}}</ref> It closely resembled living lions with a coat of yellowish-grey fur though unlike extant lions, males appear to have lacked manes.
 ''Panthera spelaea'' interacted with both [[Neanderthals]] and modern humans, who used their pelts and in the case of the latter, depicted them in artistic works.
@@ Line 26: / Line 26: @@
 Several authors regarded ''Panthera spelaea'' as a [[subspecies]] of the modern [[lion]], and therefore as ''Panthera leo spelaea''.<ref>{{cite book |author=Kurtén, B. |date=1968 |title=Pleistocene Mammals of Europe |publisher=Weidenfeld and Nicolson |location=London}}</ref><ref name=Hemmer>{{cite journal |author=Hemmer, H. |year=1974 |title=Untersuchungen zur Stammesgeschichte der Pantherkatzen (''Pantherinae'') Teil 3. Zur Artgeschichte des Löwen ''Panthera (Panthera) leo'' (Linnaeus, 1758) |journal=Veröffentlichungen der Zoologischen Staatssammlung 17 |pages=167–280 |url=https://archive.org/stream/verfentlichungen171974zool#page/178/mode/2up}}</ref><ref>{{cite journal |author=Turner, A. |year=1984 |title=Dental sex dimorphism in European lions (''Panthera leo'' L.) of the Upper Pleistocene: palaeoecological and palaeoethological implications |journal=Annales Zoologici Fennici |volume=21 |pages=1–8}}</ref><ref name=Burger/>
-One author considered the cave lion to be more closely related to the [[tiger]] based on a comparison of [[skull]] shapes, and proposed the scientific name ''Panthera tigris spelaea''.<ref>{{cite journal |author=Groiss, J. Th. |year=1996 |title=Der Höhlentiger ''Panthera tigris spelaea'' (Goldfuss) |journal=[[Neues Jahrbuch für Geologie und Paläontologie]] |issue=7 |pages=399–414 |doi=10.1127/njgpm/1996/1996/399}}</ref> Some authors regard the larger Middle Pleisto
+One author considered the cave lion to be more closely related to the [[tiger]] based on a comparison of [[skull]] shapes, and proposed the scientific name ''Panthera tigris spelaea''.<ref>{{cite journal |author=Groiss, J. Th. |year=1996 |title=Der Höhlentiger ''Panthera tigris spelaea'' (Goldfuss) |journal=[[Neues Jahrbuch für Geologie und Paläontologie]] |issue=7 |pages=399–414 |doi=10.1127/njgpm/1996/1996/399}}</ref>
 Results from morphological studies showed that it is distinct in cranial and dental [[anatomy]] to justify the [[Species|specific]] status of ''Panthera spelaea''.<ref>{{cite book |author=Spassov, N. |author2=Iliev, N. |name-list-style=amp |year=1994 |title=Proceedings of the International Symposium VI |location=Thracia Pontica |chapter=Animal remains from the submerged Late Eneolithic – early Bronze Age settlements in Sozopol (South Bulgarian Black Sea Coast) |pages=287–314}}</ref><ref>{{cite journal |author=Sotnikova, M. |author2=Nikolskiy, P. |year=2006 |title=Systematic position of the cave lion ''Panthera spelaea'' (Goldfuss) based on cranial and dental characters |journal=Quaternary International |volume=142-143 |pages=218–228 |doi=10.1016/j.quaint.2005.03.019 |bibcode=2006QuInt.142..218S |url=http://doc.rero.ch/record/13484/files/PAL_E260.pdf |access-date=22 January 2023 |archive-date=5 February 2023 |archive-url=https://web.archive.org/web/20230205061457/https://doc.rero.ch/record/13484/files/PAL_E260.pdf |url-status=live }}</ref> Results of [[Phylogenetics|phylogenetic]] studies also support this assessment.<ref>{{cite journal |last1=Christiansen |first1=P. |year=2008 |title=Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters |journal=[[Cladistics (journal)|Cladistics]] |volume=24 |issue=6 |pages=977–992 |doi=10.1111/j.1096-0031.2008.00226.x |pmid=34892880 |s2cid=84497516|doi-access=free }}</ref><ref name="Barnett2009">{{cite journal |author1=Barnett, R. |author2=Shapiro, B. |author3=Barnes, I. A. N. |author4=Ho, S. Y. |author5=Burger, J. |author6=Yamaguchi, N. |author7=Higham, T. F. |author8=Wheeler, H. |author9=Rosendahl, W. |author10=Sher, A. V. |author11=Sotnikova, M. |name-list-style=amp |year=2009 |title=Phylogeography of lions (''Panthera leo ssp.'') reveals three distinct taxa and a late Pleistocene reduction in genetic diversity |url=http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Barnett%20et%20al.%202009.pdf |journal=[[Molecular Ecology]] |volume=18 |issue=8 |pages=1668–1677 |bibcode=2009MolEc..18.1668B |doi=10.1111/j.1365-294X.2009.04134.x |pmid=19302360 |s2cid=46716748 |archive-url=https://web.archive.org/web/20120121153712/http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Barnett%20et%20al.%202009.pdf |archive-date=21 January 2012}}</ref><ref name="Barnett2016">{{cite journal |last1=Barnett |first1=R. |last2=Mendoza |first2=M. L. Z. |last3=Soares |first3=A. E. R. |last4=Ho |first4=S. Y. W. |last5=Zazula |first5=G. |last6=Yamaguchi |first6=N. |last7=Shapiro |first7=B. |last8=Kirillova |first8=I. V. |last9=Larson |first9=G. |last10=Gilbert |first10=M. T. P. |year=2016 |title=Mitogenomics of the Extinct Cave Lion, ''Panthera spelaea'' (Goldfuss, 1810), Resolve its Position within the ''Panthera'' Cats |journal=Open Quaternary |volume=2 |page=4 |doi=10.5334/oq.24 |doi-access=free |hdl-access=free |hdl=10576/22920}}</ref>
 In 2001, the subspecies ''Panthera spelaea vereshchagini'' was proposed for seven specimens found in [[Siberia]] and [[Yukon]], which have smaller skulls and teeth than the average ''P. spelaea''.<ref>{{cite journal |author1=Baryshnikov, G. F. |author2=Boeskorov, G. |name-list-style=amp |year=2001 |title=The Pleistocene cave lion, ''Panthera spelaea'' (Carnivora, Felidae) from Yakutia, Russia |journal=Cranium |volume=18 |issue=1 |pages=7–24 |url=http://natuurtijdschriften.nl/download?type=document;docid=523468 |access-date=31 December 2017 |archive-date=1 January 2018 |archive-url=https://web.archive.org/web/20180101030312/http://natuurtijdschriften.nl/download?type=document;docid=523468 |url-status=live }}</ref> Before 2020, genetic analysis using ancient DNA provided no evidence for their distinct subspecific status; DNA signatures from ''P. spelaea'' from Europe and Alaska were indistinguishable, suggesting one large [[panmictic]] population.<ref name=Barnett2009/><ref name=Stanton2020>{{cite journal |author1=Stanton, D.W. |name-list-style=amp |author2=Alberti, F. |author3=Plotnikov, V. |author4=Androsov, S. |author5=Grigoriev, S. |author6=Fedorov, S. |author7=Kosintsev, P. |author8=Nagel, D. |author9=Vartanyan, S. |author10=Barnes, I. |author11=Barnett, R. |journal=Scientific Reports |volume=10 |page=12621 |year=2020 |title=Early Pleistocene origin and extensive intra-species diversity of the extinct cave lion |issue=1 |doi=10.1038/s41598-020-69474-1 |pmid=32724178 |pmc=7387438 |bibcode=2020NatSR..1012621S |doi-access=free}}</ref> However, analysis of [[mitochondrial genome]] sequences from 31 cave lions showed that they fall into two [[Monophyletic group|monophyletic]] [[clade]]s. One lived across western Europe and the other was restricted to Beringia during the [[Pleistocene]]. For this reason, the Beringian population is considered a distinct subspecies, ''P. s. vereshchagini''.<ref name=Stanton2020 /><ref name="Boeskorov_al.2021">{{cite journal |last1=Boeskorov |first1=G. G. |last2=Plotnikov |first2=V. V. |last3=Protopopov |first3=A. V. |last4=Baryshnikov |first4=G. F. |last5=Fosse |first5=P. |last6=Dalén |first6=L. |last7=David |first7=S. W. G. |last8=Pavlov |first8=I. S. |last9=Naoki |first9=S. |last10=Alexey |first10=N. T. |name-list-style=amp |title=The preliminary analysis of Cave Lion cubs ''Panthera spelaea'' (Goldfuss, 1810) from the permafrost of Siberia |journal=[[Quaternary (journal)|Quaternary]] |volume=4 |issue=3 |page=24 |doi=10.3390/quat4030024 |doi-access=free |date=2021}}</ref>
 ===Evolution===
+Lion-like [[Panthera|pantherine]] felids first appeared in the [[Tanzania]]n [[Olduvai Gorge]] about {{mya|1.7|1.2}}. These cats dispersed into Eurasia from [[East Africa]] around the end of the [[Early Pleistocene]] and the beginning of the [[Middle Pleistocene]], giving rise to ''[[Panthera fossilis]].'' The oldest widely accepted fossils of ''P. fossilis'' in Europe date to around 700,000 years ago,<ref name=Sotnikova2014>{{cite journal |author1=Sotnikova, M.V. |author2=Foronova, I.V. |name-list-style=amp |year=2014 |title=First Asian record of ''Panthera (Leo) fossilis'' (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia |journal=Integrative Zoology |volume=9  |issue=4 |pages=517–530|doi=10.1111/1749-4877.12082 |pmid=24382145}}</ref><ref name=":0">{{Cite journal |last1=Marciszak |first1=Adrian |last2=Lipecki |first2=Grzegorz |last3=Pawłowska |first3=Kamilla |last4=Jakubowski |first4=Gwidon |last5=Ratajczak-Skrzatek |first5=Urszula |last6=Zarzecka-Szubińska |first6=Katarzyna |last7=Nadachowski |first7=Adam |date=20 December 2021 |title=The Pleistocene lion Panthera spelaea (Goldfuss, 1810) from Poland – A review |url=https://www.sciencedirect.com/science/article/pii/S1040618220308399 |url-status=live |journal=[[Quaternary International]] |series=The Quaternary of Europe and Adjacent Areas: Stratigraphical Perspectives and Tools for Correlations – SEQS-2019 |volume=605-606 |pages=213–240 |bibcode=2021QuInt.605..213M |doi=10.1016/j.quaint.2020.12.018 |issn=1040-6182 |archive-url=https://web.archive.org/web/20240518071941/https://www.sciencedirect.com/science/article/abs/pii/S1040618220308399 |archive-date=18 May 2024 |access-date=22 March 2024 |via=Elsevier Science Direct}}</ref><ref name=":2" /><ref name="Ianucci_al.2024">{{cite journal |last1=Iannucci |first1=A. |last2=Mecozzi |first2=B. |last3=Pineda |first3=A. |last4=Raffaele |first4=S. |last5=Carpentieri |first5=M. |last6=Rabinovich |first6=R. |last7=Moncel |first7=M.-H. |title=Early occurrence of lion (Panthera spelaea) at the Middle Pleistocene Acheulean site of Notarchirico (MIS 16, Italy) |journal=[[Journal of Quaternary Science|Quaternary Science]] |doi=10.1002/jqs.3639 |date=2024-06-24 |volume=39 |issue=5 |pages=683–690 |issn=0267-8179 |doi-access=free|bibcode=2024JQS....39..683I }}</ref> with possible older fossils from [[Western Siberia]] dating to the late Early Pleistocene.<ref>{{Cite journal |last=Sotnikova |first=Marina V. |last2=Foronova |first2=Irina V. |date=August 2014 |title=First Asian record of Panthera (Leo) fossilis (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia |url=https://onlinelibrary.wiley.com/doi/10.1111/1749-4877.12082 |journal=Integrative Zoology |volume=9 |issue=4 |pages=517–530 |doi=10.1111/1749-4877.12082 |issn=1749-4877}}</ref> Different authors considered ''Panthera fossils'' as either a distinct species ancestral to ''P. spelaea'',<ref name=":02">{{Cite journal |last1=Sabo |first1=Martin |last2=Tomašových |first2=Adam |last3=Gullár |first3=Juraj |date=August 2022 |title=Geographic and temporal variability in Pleistocene lion-like felids: Implications for their evolution and taxonomy |url=https://palaeo-electronica.org/content/2022/3681-lion-cranial-variability |journal=Palaeontologia Electronica |volume=25 |issue=2 |pages=1–27 |doi=10.26879/1175 |issn=1094-8074 |s2cid=251855356 |doi-access=free}}</ref> or as a subspecies of ''P. spelaea''.<ref name="Ianucci_al.2024" /><ref name=":1">{{Cite journal |last1=Marciszak |first1=Adrian |last2=Ivanoff |first2=Dmitry V. |last3=Semenov |first3=Yuriy A. |last4=Talamo |first4=Sahra |last5=Ridush |first5=Bogdan |last6=Stupak |first6=Alina |last7=Yanish |first7=Yevheniia |last8=Kovalchuk |first8=Oleksandr |date=March 2023 |title=The Quaternary lions of Ukraine and a trend of decreasing size in Panthera spelaea |journal=Journal of Mammalian Evolution |volume=30 |issue=1 |pages=109–135 |doi=10.1007/s10914-022-09635-3 |issn=1064-7554 |doi-access=free}}</ref> Recent nuclear genomic evidence suggest that interbreeding between modern lions and all Eurasian fossil lions took place up until 500,000 years ago, but by 470,000 years ago, no subsequent interbreeding between the two lineages occurred.<ref name=Sotnikova2014/><ref name=Burger/><ref name=Manuel_al2020/>
-[[File:18 PanneauDesLions(PartieDroite)BisonsPoursuivisParDesLions.jpg|thumb|Cave lions and [[bison]] depicted in the [[Chauvet Cave]], [[France]]<ref name=Chauvet1996>{{cite book |author=Chauvet, J.-M. |author2=Brunel, D. E. |author3=Hillaire, C. |name-list-style=amp |year=1996 |title=Dawn of Art: The Chauvet Cave. The oldest known paintings in the world |url=https://archive.org/details/dawnofartchauvet0000chau |url-access=registration |location=New York |publisher=Harry N. Abrams}}</ref>]]
-Lion-like [[Panthera|pantherine]] felids first appeared in the [[Tanzania]]n [[Olduvai Gorge]] about {{mya|1.7|1.2}}. These cats dispersed to Europe from [[East Africa]] in the first half of the Middle Pleistocene, giving rise to ''[[Panthera fossilis]].'' The oldest widely accepted fossils of ''P. fossilis'' in Europe date to around 700,000 years ago.<ref name=Sotnikova2014>{{cite journal |author1=Sotnikova, M.V. |author2=Foronova, I.V. |name-list-style=amp |year=2014 |title=First Asian record of ''Panthera (Leo) fossilis'' (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia |journal=Integrative Zoology |volume=9  |issue=4 |pages=517–530|doi=10.1111/1749-4877.12082 |pmid=24382145}}</ref><ref name=":0">{{Cite journal |last1=Marciszak |first1=Adrian |last2=Lipecki |first2=Grzegorz |last3=Pawłowska |first3=Kamilla |last4=Jakubowski |first4=Gwidon |last5=Ratajczak-Skrzatek |first5=Urszula |last6=Zarzecka-Szubińska |first6=Katarzyna |last7=Nadachowski |first7=Adam |date=20 December 2021 |title=The Pleistocene lion Panthera spelaea (Goldfuss, 1810) from Poland – A review |url=https://www.sciencedirect.com/science/article/pii/S1040618220308399 |url-status=live |journal=[[Quaternary International]] |series=The Quaternary of Europe and Adjacent Areas: Stratigraphical Perspectives and Tools for Correlations – SEQS-2019 |volume=605-606 |pages=213–240 |bibcode=2021QuInt.605..213M |doi=10.1016/j.quaint.2020.12.018 |issn=1040-6182 |archive-url=https://web.archive.org/web/20240518071941/https://www.sciencedirect.com/science/article/abs/pii/S1040618220308399 |archive-date=18 May 2024 |access-date=22 March 2024 |via=Elsevier Science Direct}}</ref><ref name=":2" /><ref name="Ianucci_al.2024">{{cite journal |last1=Iannucci |first1=A. |last2=Mecozzi |first2=B. |last3=Pineda |first3=A. |last4=Raffaele |first4=S. |last5=Carpentieri |first5=M. |last6=Rabinovich |first6=R. |last7=Moncel |first7=M.-H. |title=Early occurrence of lion (Panthera spelaea) at the Middle Pleistocene Acheulean site of Notarchirico (MIS 16, Italy) |journal=[[Journal of Quaternary Science|Quaternary Science]] |doi=10.1002/jqs.3639 |date=2024-06-24 |volume=39 |issue=5 |pages=683–690 |issn=0267-8179 |doi-access=free}}</ref> Different authors considered ''Panthera fossils'' as either a distinct species ancestral to ''P. spelaea'',<ref name=":02">{{Cite journal |last1=Sabo |first1=Martin |last2=Tomašových |first2=Adam |last3=Gullár |first3=Juraj |date=August 2022 |title=Geographic and temporal variability in Pleistocene lion-like felids: Implications for their evolution and taxonomy |url=https://palaeo-electronica.org/content/2022/3681-lion-cranial-variability |journal=Palaeontologia Electronica |language=English |volume=25 |issue=2 |pages=1–27 |doi=10.26879/1175 |issn=1094-8074 |s2cid=251855356 |doi-access=free}}</ref> or as a subspecies of ''P. spelaea''.<ref name="Ianucci_al.2024" /><ref name=":1">{{Cite journal |last1=Marciszak |first1=Adrian |last2=Ivanoff |first2=Dmitry V. |last3=Semenov |first3=Yuriy A. |last4=Talamo |first4=Sahra |last5=Ridush |first5=Bogdan |last6=Stupak |first6=Alina |last7=Yanish |first7=Yevheniia |last8=Kovalchuk |first8=Oleksandr |date=March 2023 |title=The Quaternary lions of Ukraine and a trend of decreasing size in Panthera spelaea |journal=Journal of Mammalian Evolution |language=en |volume=30 |issue=1 |pages=109–135 |doi=10.1007/s10914-022-09635-3 |issn=1064-7554 |doi-access=free}}</ref> Recent nuclear genomic evidence suggest that interbreeding between modern lions and all Eurasian fossil lions took place up until 500,000 years ago, but by 470,000 years ago, no subsequent interbreeding between the two lineages occurred.<ref name=Sotnikova2014/><ref name=Burger/><ref name=Manuel_al2020/>
 The following cladogram shows the genetic relationship between ''P. spelaea'' and other pantherine cats.<ref name="Barnett2016" />{{clade| style=font-size:100%; line-height:100%
@@ Line 55: / Line 54: @@
        |1={{clade
         |1=''[[Panthera leo leo]]''<span style="{{MirrorH}}">[[File:W. Gornig - P. leo leo ♂.png|70 px]]</span> (Asian, North and West African lions)
-        |2=''[[Panthera leo melanochaita]]''<span style="{{MirrorH}}">[[File:W. Gornig - P. leo melanochaita ♂.png|70 px]]</span>}} (Southern and East African lions)
+        |2=''[[Panthera leo melanochaita]]''<span style="{{MirrorH}}">[[File:W. Gornig - P. leo melanochaita ♂.png|70 px]]</span> (Southern and East African lions)}}
-       |2={{extinct}}'''''Panthera spelaea'''''<span style="{{MirrorH}}">[[File:W. Gornig - P. spelaea spelaea.png|90 px]]</span> }} }} }} }} }} }} }} }}Specimens intermediate between ''P. fossilis'' and Late Pleistocene ''P. spelaea'' are referred to as the subspecies ''P. s. intermedia''.<ref name=":0" /> The transition from ''P. fossilis'' to Late Pleistocene ''P. spelaea'' involved significant changes in skull and tooth morphology.<ref>{{Cite journal |last=Persico |first=Davide |date=June 2021 |title=First fossil record of cave lion (Panthera (Leo) spelaea intermedia) from alluvial deposits of the Po River in northern Italy |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618221000902 |journal=Quaternary International |language=en |volume=586 |pages=14–23 |doi=10.1016/j.quaint.2021.02.029|bibcode=2021QuInt.586...14P }}</ref> [[Mitochondrial DNA]] sequence data from fossil lion remains show that the [[American lion]] represents a [[sister group]] of Late Pleistocene ''P. spelaea'', and likely arose when an early ''P. spelaea'' population became isolated south of the [[Cordilleran Ice Sheet]]. Initially this was suggested to be around 340,000 years ago,<ref name="Barnett2009" /> but later studies suggested that the split between the two species was probably younger, around 165,000 years ago, consistent with the late first appearance of ''P. spelaea'' in Eastern [[Beringia]] (now Alaska and adjacent regions) during the [[Illinoian (stage)|Illinoian]] (around 190-130,000 years ago).<ref>{{Cite journal |last1=Salis |first1=Alexander T. |last2=Bray |first2=Sarah C. E. |last3=Lee |first3=Michael S. Y. |last4=Heiniger |first4=Holly |last5=Barnett |first5=Ross |last6=Burns |first6=James A. |last7=Doronichev |first7=Vladimir |last8=Fedje |first8=Daryl |last9=Golovanova |first9=Liubov |last10=Harington |first10=C. Richard |last11=Hockett |first11=Bryan |last12=Kosintsev |first12=Pavel |last13=Lai |first13=Xulong |last14=Mackie |first14=Quentin |last15=Vasiliev |first15=Sergei |date=December 2022 |title=Lions and brown bears colonized North America in multiple synchronous waves of dispersal across the Bering Land Bridge |url=https://onlinelibrary.wiley.com/doi/10.1111/mec.16267 |journal=Molecular Ecology |language=en |volume=31 |issue=24 |pages=6407–6421 |doi=10.1111/mec.16267 |bibcode=2022MolEc..31.6407S |issn=0962-1083|hdl=11343/299180 |hdl-access=free }}</ref>
+       |2={{extinct}}'''''Panthera spelaea'''''<span style="{{MirrorH}}">[[File:W. Gornig - P. spelaea spelaea.png|90 px]]</span> }} }} }} }} }} }} }} }}Specimens intermediate between ''P. fossilis'' and Late Pleistocene ''P. spelaea'' are referred to as the subspecies ''P. s. intermedia''.<ref name=":0" /> The transition from ''P. fossilis'' to Late Pleistocene ''P. spelaea'' involved significant changes in skull and tooth morphology.<ref>{{Cite journal |last=Persico |first=Davide |date=June 2021 |title=First fossil record of cave lion (Panthera (Leo) spelaea intermedia) from alluvial deposits of the Po River in northern Italy |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618221000902 |journal=Quaternary International |volume=586 |pages=14–23 |doi=10.1016/j.quaint.2021.02.029|bibcode=2021QuInt.586...14P }}</ref> [[Mitochondrial DNA]] sequence data from fossil lion remains show that the [[American lion]] represents a [[sister group]] of Late Pleistocene ''P. spelaea'', and likely arose when an early ''P. spelaea'' population became isolated south of the [[Cordilleran Ice Sheet]]. Initially this was suggested to be around 340,000 years ago,<ref name="Barnett2009" /> but later studies suggested that the split between the two species was probably younger, around 165,000 years ago, consistent with the late first appearance of ''P. spelaea'' in Eastern [[Beringia]] (now Alaska and adjacent regions) during the [[Illinoian (stage)|Illinoian]] (around 190-130,000 years ago).<ref>{{Cite journal |last1=Salis |first1=Alexander T. |last2=Bray |first2=Sarah C. E. |last3=Lee |first3=Michael S. Y. |last4=Heiniger |first4=Holly |last5=Barnett |first5=Ross |last6=Burns |first6=James A. |last7=Doronichev |first7=Vladimir |last8=Fedje |first8=Daryl |last9=Golovanova |first9=Liubov |last10=Harington |first10=C. Richard |last11=Hockett |first11=Bryan |last12=Kosintsev |first12=Pavel |last13=Lai |first13=Xulong |last14=Mackie |first14=Quentin |last15=Vasiliev |first15=Sergei |date=December 2022 |title=Lions and brown bears colonized North America in multiple synchronous waves of dispersal across the Bering Land Bridge |url=https://onlinelibrary.wiley.com/doi/10.1111/mec.16267 |journal=Molecular Ecology |volume=31 |issue=24 |pages=6407–6421 |doi=10.1111/mec.16267 |pmid=34748674 |bibcode=2022MolEc..31.6407S |issn=0962-1083|hdl=11343/299180 |hdl-access=free }}</ref>
 ==Characteristics==
+[[File:18 PanneauDesLions(PartieDroite)BisonsPoursuivisParDesLions.jpg|thumb|Cave lions and [[bison]] depicted in the [[Chauvet Cave]], [[France]]<ref name="Chauvet1996">{{cite book |author=Chauvet, J.-M. |author2=Brunel, D. E. |author3=Hillaire, C. |name-list-style=amp |year=1996 |title=Dawn of Art: The Chauvet Cave. The oldest known paintings in the world |url=https://archive.org/details/dawnofartchauvet0000chau |url-access=registration |location=New York |publisher=Harry N. Abrams}}</ref>]]
-[[File:Lascaux-diverticule-félins.jpg|thumb|Cave lions, Chamber of Felines, [[Lascaux]] caves]]
 Carvings and [[cave paintings]] of cave lions, which were discovered in the [[Lascaux]] and [[Chauvet Cave]]s in France, were dated to 15,000 to 17,000 years old.<ref name=Chauvet1996 /><ref>{{cite book |author=Leroi-Gourhan, A. |author2=Allain, J. |date=1979 |title=Lascaux inconnu. XXIIe supplement à "Gallia Préhistoire" |location=Paris}}</ref> A drawing in the Chauvet cave depicts two cave lions walking together. The one in the foreground is slightly smaller than the one in the background, which has been drawn with a [[scrotum]] and without a mane.<ref name=Yamaguchietal2004>{{cite journal |last1=Yamaguchi |first1=N. | last2=Cooper |first2=A. |last3=Werdelin |first3=L. |last4= MacDonald |first4=D. W. |date=2004 |title=Evolution of the mane and group-living in the lion (''Panthera leo''): a review |journal=Journal of Zoology |volume=263 |issue=4 |pages=329–342 |doi=10.1017/S0952836904005242}}</ref> Such cave paintings suggest that male cave lions completely lacked manes, or at most had very small manes.<ref name=Stuart2011/>
+[[File:Panthera spelaea size comparison.jpg|thumb|Size comparison of various specimens attributed to the ''P. spelaea''/''fossilis'' lineage. 1 =''P.'' (''spelaea'') ''fossilis'' male (M) from Château, France MIS ([[Marine Isotope Stage]]) 15 ~ 600,000 years ago 2= ''P. spelaea'' from Niedźwiedzia Cave, Poland, MIS 5 (130-80,000 years ago). 3 = M from Siegsdorf, Germany MIS 3 (57-29,000 years ago) 4= female (F) from Zandobbio, Italy, [[Last Interglacial]] (130-115,000 years ago) 5=  F from Kryshtaleva Cave, Ukraine (21-22,000 years ago). Scale bar = 10 cm]]
-[[File:W. Gornig - P. spelaea spelaea.png|thumb|left|Life restoration]]Some individuals of ''P. spelaea'' were considerably larger than modern lions. The skeleton of an adult male found in 1985 near [[Siegsdorf]] in Germany had a shoulder height of around {{convert|1.2|m|ftin|abbr=on}} and a head-body length of {{convert|2|m|ftin|abbr=on}} without the tail, similar in size to large modern lions.<ref>{{cite journal|first1=S.T.|last1=Hussain|first2=H.|last2=Floss|year=2015|title=Sharing the world with mammoths, cave lions and other beings: linking animal-human interactions and the Aurignacian "belief world"|journal=Quartär|volume=65|pages=85–120|doi=10.7485/QU62_4}}</ref> Similarly, footprints attributed to a male cave lion  measured  {{convert|15|cm|in|0|abbr=on}} across. The heaviest  ''Panthera spelaea'' was estimated to weigh {{convert|339|kg|lb|abbr=on}}.<ref>{{cite journal |last=Sherani |first=Shaheer |title=A new specimen-dependent method of estimating felid body mass |journal=PeerJ Preprints |year=2016 |page=16 |doi=10.7287/peerj.preprints.2327v2 |url=https://peerj.com/preprints/2327.pdf |doi-access=free |access-date=15 March 2021 |archive-date=20 September 2022 |archive-url=https://web.archive.org/web/20220920173645/https://peerj.com/preprints/2327.pdf |url-status=dead }}</ref> This shows that some individuals of ''P. spelaea'' would have been up to or over 12% larger than modern lions, but still smaller than the earlier ''Panthera fossilis'' or the [[American lion]] (''P. atrox'').<ref name="Koenigswald02">{{Cite book |last=Koenigswald |first=W. v. |title=Lebendige Eiszeit: Klima und Tierwelt im Wandel |publisher=Theiss |year=2002 |isbn=978-3-8062-1734-6 |location=Stuttgart |language=de}}</ref><ref>{{cite book |author=Koenigswald, W. v. |title=Lebendige Eiszeit |date=2002 |publisher=Theiss-Wissenschaftliche Buchgemeinschaft |isbn=3-8062-1734-3 |location=Darmstadt}}</ref> Early members of the cave lion lineage (including those assigned ''Panthera fossils'') were considerably larger than individuals of ''P. spelaea'' from the [[Last Glacial Period]], which were comparable in size to modern lions, with the species showing a progressive size reduction over the course of the Last Glacial Period up until its extinction, with the last ''P. spelaea'' populations comparable in size to small-sized modern lions.<ref name=":1" />
+Early members of the cave lion lineage (including those assigned ''Panthera fossilis'') during the Middle Pleistocene were considerably larger than individuals of ''P. spelaea'' from the [[Last Glacial Period]] and modern lions, with some of these individuals having an estimated length of {{Convert|2.5-2.9|m|ft}}, shoulder height of {{Convert|1.4-1.5|m|ft}} and body mass of {{Convert|400-500|kg}}, respectively, making them among the largest cats to have ever lived. The Late Pleistocene ''Panthera spelaea spelaea'' was noticeably smaller though still large relative to living cats, with an estimated length and  shoulder height of {{Convert|2-2.1|m|ft}} and {{Convert|1.1-1.2|m|ft}}, respectively, The species showed a progressive size reduction over the course of the Last Glacial Period up until its extinction, with the last ''P. spelaea'' populations comparable in size to small-sized modern lions, with a body mass of only {{Convert|70-90|kg}} and a body length and shoulder height of only {{Convert|1.2-1.3|m|ft}} and {{Convert|70-75|cm|ft}} respectively.<ref name=":1" /><ref>{{Cite journal |last=Marciszak |first=Adrian |last2=Gornig |first2=Wiktoria |date=September 2024 |title=From giant to dwarf: A trend of decreasing size in Panthera spelaea (Goldfuss, 1810) and its likely implications |url=https://linkinghub.elsevier.com/retrieve/pii/S2950475924000078 |journal=Earth History and Biodiversity |language=en |volume=1 |pages=100007 |doi=10.1016/j.hisbio.2024.100007|doi-access=free }}</ref>
 ''P. spelaea'' had a relatively longer and narrower muzzle compared to that of the extant lion. Despite this, the two species do not exhibit major differences in morphology.<ref name="Stuart2011" /> Like modern lions, females were smaller than males.<ref>{{cite journal |author=Diedrich, C. G. |year=2011 |title=Late Pleistocene ''Panthera leo spelaea'' (Goldfuss, 1810) skeletons from the Czech Republic (central Europe); their pathological cranial features and injuries resulting from intraspecific fights, conflicts with hyenas, and attacks on cave bears |journal=Bulletin of Geosciences |volume=86 |issue=4 |pages=817–840 |doi=10.3140/bull.geosci.1263 |doi-access=free }}</ref>
@@ Line 68: / Line 68: @@
 ==Distribution and habitat==
-[[File:Ice age fauna of northern Spain - Mauricio Antón.jpg|thumb|Cave lions and other Ice Age fauna in northern Spain, by [[Mauricio Antón]]]]
+[[File:Ice age fauna of northern Spain - Mauricio Antón.jpg|thumb|Cave lions feeding on [[reindeer]] in a [[mammoth steppe]] landscape in northern Spain also including [[woolly rhinoceros]], [[woolly mammoth]] and [[wild horse]]. Art by [[Mauricio Antón]]]]
-During the Late Pleistocene, ''P. spelaea'' formed a contiguous population across the [[mammoth steppe]], from Western Europe to northwest North America.<ref name="Stuart2011" /><ref>{{cite journal |author=Ersmark, E. |author2=Orlando, L. |author3=Sandoval-Castellanos, E. |author4=Barnes, I. |author5=Barnett, R. |author6=Stuart, A. |author7=Lister, A. |author8=Dalén, L. |year=2015 |title=Population Demography and Genetic Diversity in the Pleistocene Cave Lion |journal=[[Open Quaternary]] |volume=1 |issue=1 |pages=Art. 4 |doi=10.5334/oq.aa |doi-access=free}}</ref> It was widely distributed in the [[Iberian Peninsula]],<ref>{{Cite journal |last1=Álvarez-Lao |first1=D. J. |last2=Álvarez-Vena |first2=A. |last3=Ballesteros |first3=D. |last4=García |first4=N. |last5=Laplana |first5=C. |date=2020 |title=A cave lion (''Panthera spelaea'') skeleton from Torca del León (NW Iberia): Micromammals indicate a temperate and forest environment corresponding to GI-11 (MIS 3) |journal=[[Quaternary Science Reviews]] |volume=229 |page=106123 |bibcode=2020QSRv..22906123A |doi=10.1016/j.quascirev.2019.106123 |hdl-access=free |hdl=10651/54312}}</ref> [[Italian Peninsula]],<ref>Iannucci, Alessio, Mecozzi, Beniamino, Pineda, Antonio, Sardella, Raffaele, Carpentieri, Marco, Rabinovich, Rivka, and Moncel Marie-Helene. "Early Occurence of (''Panthera spelaea'') at the Middle Pleistocene Acheulean site of Notarchirico (MIS 16, Italy). ''Journal of Quarternary Sciences'' 39:3 (2024): 4.</ref> [[Southeast Europe]],<ref>{{Cite journal |last1=Smith |first1=Geoff M. |last2=Spasov |first2=Rosen |last3=Martisius |first3=Naomi L. |last4=Sinet-Mathiot |first4=Virginie |last5=Aldeias |first5=Vera |last6=Rezek |first6=Zeljko |last7=Ruebens |first7=Karen |last8=Pederzani |first8=Sarah |last9=McPherron |first9=Shannon P. |last10=Sirakova |first10=Svoboda |last11=Sirakov |first11=Nikolay |last12=Tsanova |first12=Tsenka |last13=Hublin |first13=Jean-Jacques |date=December 2021 |title=Subsistence behavior during the Initial Upper Paleolithic in Europe: Site use, dietary practice, and carnivore exploitation at Bacho Kiro Cave (Bulgaria) |url=https://linkinghub.elsevier.com/retrieve/pii/S0047248421001263 |journal=[[Journal of Human Evolution]] |language=en |volume=161 |pages=103074 |doi=10.1016/j.jhevol.2021.103074 |pmid=34628301 |bibcode=2021JHumE.16103074S |access-date=22 March 2024 |via=Elsevier Science Direct}}</ref> [[Great Britain]],<ref name="Stuart2011" /> [[Central Europe]],<ref>{{Cite journal |last1=Sabol |first1=Martin |last2=Döppes |first2=Doris |date=January 2020 |title=The first radiocarbon dating of a cave lion fossil from the Slovakian Western Carpathians |url=https://www.researchgate.net/publication/338937589 |journal=Acta Geologica Slovaca |volume=11 |issue=1 |pages=11–14 |via=ResearchGate}}</ref><ref>{{Cite journal |last1=Diedrich |first1=Cajus G. |last2=Rathgeber |first2=Thomas |date=12 October 2011 |title=Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss 1810) skeleton remains of the Upper Rhine Valley (SW Germany) and contributions to their sexual dimorphism, taphonomy and habitus |url=http://www.tandfonline.com/doi/abs/10.1080/08912963.2010.549943 |journal=[[Historical Biology]] |language=en |volume=24 |issue=1 |pages=1–28 |doi=10.1080/08912963.2010.549943 |issn=0891-2963 |access-date=24 March 2024 |via=Taylor and Francis Online |archive-date=18 May 2024 |archive-url=https://web.archive.org/web/20240518071934/https://www.tandfonline.com/doi/abs/10.1080/08912963.2010.549943 |url-status=live }}</ref><ref>{{Cite journal |last1=Marciszak |first1=Adrian |last2=Schouwenburg |first2=Charles |last3=Gornig |first3=Wiktoria |last4=Lipecki |first4=Grzegorz |last5=Mackiewicz |first5=Paweł |date=1 November 2019 |title=Morphometric comparison of Panthera spelaea (Goldfuss, 1810) from Poland with the lion remains from Eurasia over the last 700 ka |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119303907 |journal=[[Quaternary Science Reviews]] |language=en |volume=223 |pages=105950 |doi=10.1016/j.quascirev.2019.105950 |bibcode=2019QSRv..22305950M |access-date=24 March 2024 |via=Elsevier Science Direct |archive-date=29 March 2023 |archive-url=https://web.archive.org/web/20230329081534/https://linkinghub.elsevier.com/retrieve/pii/S0277379119303907 |url-status=live }}</ref> the [[East European Plain]],<ref name="Stuart2011" /> the [[Ural Mountains]],<ref>{{Cite journal |last1=Gimranov |first1=Dmitry |last2=Kosintsev |first2=Pavel |date=30 April 2020 |title=Quaternary large mammals from the Imanay Cave |url=https://www.sciencedirect.com/science/article/pii/S104061822030015X |journal=[[Quaternary International]] |series=Quaternary Stratigraphy and Karst & Cave Sediments: the INQUA-SEQS 2018 Meeting |volume=546 |pages=125–134 |doi=10.1016/j.quaint.2020.01.014 |bibcode=2020QuInt.546..125G |issn=1040-6182 |access-date=22 March 2024 |via=Elsevier Science Direct |archive-date=18 May 2024 |archive-url=https://web.archive.org/web/20240518071940/https://www.sciencedirect.com/science/article/abs/pii/S104061822030015X |url-status=live }}</ref> most of [[North Asia|Northern Asia]] (ranging as far south as [[Northeast China]]<ref>{{Cite journal |last=Sherani |first=Shaheer |last2=Perng |first2=Liongvi |last3=Sherani |first3=Maryam |date=2023-06-03 |title=Evidence of cave lion ( Panthera spelaea ) from Pleistocene Northeast China |url=https://www.tandfonline.com/doi/full/10.1080/08912963.2022.2071711 |journal=Historical Biology |language=en |volume=35 |issue=6 |pages=988–996 |doi=10.1080/08912963.2022.2071711 |issn=0891-2963}}</ref>), and across the [[Bering land bridge]] into [[Alaska]] and [[Yukon]].<ref name="Stuart2011" /> The cave lion had a wide elevation range as well; in Slovakia, [[skull]], [[femur]] and [[pelvis]] remains were excavated in ten [[Karst]] caves in hilly and montane areas at elevations from {{cvt|240|to|1133|m}}.<ref>{{cite journal |author=Sabol, M. |year=2011 |title=A record of Pleistocene lion-like felids in the territory of Slovakia |journal=Quaternaire |series=Hors-série |issue=4 |pages=215−228}}</ref>
+During the [[Last Glacial Period]], ''P. spelaea'' formed a contiguous population across the [[mammoth steppe]], from Western Europe to northwest North America.<ref name="Stuart2011" /><ref>{{cite journal |author=Ersmark, E. |author2=Orlando, L. |author3=Sandoval-Castellanos, E. |author4=Barnes, I. |author5=Barnett, R. |author6=Stuart, A. |author7=Lister, A. |author8=Dalén, L. |year=2015 |title=Population Demography and Genetic Diversity in the Pleistocene Cave Lion |journal=[[Open Quaternary]] |volume=1 |issue=1 |pages=Art. 4 |doi=10.5334/oq.aa |doi-access=free}}</ref> It was widely distributed in the [[Iberian Peninsula]],<ref>{{Cite journal |last1=Álvarez-Lao |first1=D. J. |last2=Álvarez-Vena |first2=A. |last3=Ballesteros |first3=D. |last4=García |first4=N. |last5=Laplana |first5=C. |date=2020 |title=A cave lion (''Panthera spelaea'') skeleton from Torca del León (NW Iberia): Micromammals indicate a temperate and forest environment corresponding to GI-11 (MIS 3) |journal=[[Quaternary Science Reviews]] |volume=229 |page=106123 |bibcode=2020QSRv..22906123A |doi=10.1016/j.quascirev.2019.106123 |hdl-access=free |hdl=10651/54312}}</ref> [[Italian Peninsula]],<ref>Iannucci, Alessio, Mecozzi, Beniamino, Pineda, Antonio, Sardella, Raffaele, Carpentieri, Marco, Rabinovich, Rivka, and Moncel Marie-Helene. "Early Occurrence of lion (''Panthera spelaea'') at the Middle Pleistocene Acheulean site of Notarchirico (MIS 16, Italy). ''Journal of Quarternary Sciences'' 39:3 (2024): 4.</ref> [[Southeast Europe]],<ref>{{Cite journal |last1=Smith |first1=Geoff M. |last2=Spasov |first2=Rosen |last3=Martisius |first3=Naomi L. |last4=Sinet-Mathiot |first4=Virginie |last5=Aldeias |first5=Vera |last6=Rezek |first6=Zeljko |last7=Ruebens |first7=Karen |last8=Pederzani |first8=Sarah |last9=McPherron |first9=Shannon P. |last10=Sirakova |first10=Svoboda |last11=Sirakov |first11=Nikolay |last12=Tsanova |first12=Tsenka |last13=Hublin |first13=Jean-Jacques |date=December 2021 |title=Subsistence behavior during the Initial Upper Paleolithic in Europe: Site use, dietary practice, and carnivore exploitation at Bacho Kiro Cave (Bulgaria) |url=https://linkinghub.elsevier.com/retrieve/pii/S0047248421001263 |journal=[[Journal of Human Evolution]] |volume=161 |pages=103074 |doi=10.1016/j.jhevol.2021.103074 |pmid=34628301 |bibcode=2021JHumE.16103074S |access-date=22 March 2024 |via=Elsevier Science Direct}}</ref> [[Great Britain]],<ref name="Stuart2011" /> [[Central Europe]],<ref>{{Cite journal |last1=Sabol |first1=Martin |last2=Döppes |first2=Doris |date=January 2020 |title=The first radiocarbon dating of a cave lion fossil from the Slovakian Western Carpathians |url=https://www.researchgate.net/publication/338937589 |journal=Acta Geologica Slovaca |volume=11 |issue=1 |pages=11–14 |via=ResearchGate}}</ref><ref>{{Cite journal |last1=Diedrich |first1=Cajus G. |last2=Rathgeber |first2=Thomas |date=12 October 2011 |title=Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss 1810) skeleton remains of the Upper Rhine Valley (SW Germany) and contributions to their sexual dimorphism, taphonomy and habitus |url=http://www.tandfonline.com/doi/abs/10.1080/08912963.2010.549943 |journal=[[Historical Biology]] |volume=24 |issue=1 |pages=1–28 |doi=10.1080/08912963.2010.549943 |issn=0891-2963 |access-date=24 March 2024 |via=Taylor and Francis Online |archive-date=18 May 2024 |archive-url=https://web.archive.org/web/20240518071934/https://www.tandfonline.com/doi/abs/10.1080/08912963.2010.549943 |url-status=live }}</ref><ref>{{Cite journal |last1=Marciszak |first1=Adrian |last2=Schouwenburg |first2=Charles |last3=Gornig |first3=Wiktoria |last4=Lipecki |first4=Grzegorz |last5=Mackiewicz |first5=Paweł |date=1 November 2019 |title=Morphometric comparison of Panthera spelaea (Goldfuss, 1810) from Poland with the lion remains from Eurasia over the last 700 ka |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119303907 |journal=[[Quaternary Science Reviews]] |volume=223 |pages=105950 |doi=10.1016/j.quascirev.2019.105950 |bibcode=2019QSRv..22305950M |access-date=24 March 2024 |via=Elsevier Science Direct |archive-date=29 March 2023 |archive-url=https://web.archive.org/web/20230329081534/https://linkinghub.elsevier.com/retrieve/pii/S0277379119303907 |url-status=live }}</ref> the [[East European Plain]],<ref name="Stuart2011" /> the [[Ural Mountains]],<ref>{{Cite journal |last1=Gimranov |first1=Dmitry |last2=Kosintsev |first2=Pavel |date=30 April 2020 |title=Quaternary large mammals from the Imanay Cave |url=https://www.sciencedirect.com/science/article/pii/S104061822030015X |journal=[[Quaternary International]] |series=Quaternary Stratigraphy and Karst & Cave Sediments: the INQUA-SEQS 2018 Meeting |volume=546 |pages=125–134 |doi=10.1016/j.quaint.2020.01.014 |bibcode=2020QuInt.546..125G |issn=1040-6182 |access-date=22 March 2024 |via=Elsevier Science Direct |archive-date=18 May 2024 |archive-url=https://web.archive.org/web/20240518071940/https://www.sciencedirect.com/science/article/abs/pii/S104061822030015X |url-status=live }}</ref> most of [[Northeast Asia]] (ranging as far south as [[Northeast China]]<ref>{{Cite journal |last1=Sherani |first1=Shaheer |last2=Perng |first2=Liongvi |last3=Sherani |first3=Maryam |date=2023-06-03 |title=Evidence of cave lion (''Panthera spelaea'') from Pleistocene Northeast China |url=https://www.tandfonline.com/doi/full/10.1080/08912963.2022.2071711 |journal=Historical Biology |volume=35 |issue=6 |pages=988–996 |doi=10.1080/08912963.2022.2071711 |bibcode=2023HBio...35..988S |issn=0891-2963}}</ref> and possibly the [[Korean peninsula]]<ref>{{Cite journal|last1=Puzachenko |first1=A. Yu. |last2=Argant |first2=A. |last3=Baryshnikov |first3=G. F. |last4=Boeskorov |first4=G. G. |last5=Kirillova |first5=I. V. |last6=Klimovsky |first6=A. I. |last7=Kosintsev |first7=P. A. |last8=Marciszak |first8=A. |last9=Plotnikov |first9=V. V. |last10=Sabol |first10=M. |last11=Vasiliev |first11=S. K. |year=2024 |title=Distribution history of the cave lion (''Panthera spelaea'' (Goldfuss, 1810)) |journal=Earth History and Biodiversity |volume=1 |at=100006 |doi=10.1016/j.hisbio.2024.100006 |doi-access=free }}</ref>), and across the [[Bering land bridge]] into [[Alaska]] and [[Yukon]].<ref name="Stuart2011" /> The cave lion had a wide elevation range, with finds extending up over {{Convert|2000|m|ft}} above sea level in the European Alps and in [[Buryatia]] in Northern Asia, though they probably did not occupy mountainous habitats all-year round.<ref>{{Cite journal |last=Sabol |first=Martin |last2=Puzachenko |first2=Andrey Yu. |date=September 2024 |title=Distribution of cave lions (Panthera spelaea ssp.) in mountain areas |url=https://linkinghub.elsevier.com/retrieve/pii/S2950475924000091 |journal=Earth History and Biodiversity |language=en |volume=1 |pages=100009 |doi=10.1016/j.hisbio.2024.100009|doi-access=free }}</ref> The cave lion probably inhabited predominantly open habitats such as [[steppe]] and grasslands although it would have also have occurred in open woodlands as well.<ref name="Stuart2011" /> It may have sought out hibernating bears in montane caves as a food source during the winter.<ref>{{Cite journal |last1=Sabol |first1=M. |last2=Gullár |first2=J. |last3=Horvát |first3=J. |date=2018 |title=Montane record of the late Pleistocene ''Panthera spelaea'' (Goldfuss, 1810) from the Západné Tatry Mountains (northern Slovakia) |journal=[[Journal of Vertebrate Paleontology]] |volume=38 |issue=3 |pages=e1467921 |doi=10.1080/02724634.2018.1467921 |doi-access=free|bibcode=2018JVPal..38E7921S }}</ref> While during the Last Glacial Period it was often associated with cold environments, the species also inhabited temperate environments,<ref>{{Cite journal |last1=Álvarez-Lao |first1=Diego J. |last2=Álvarez-Vena |first2=Adrián |last3=Ballesteros |first3=Daniel |last4=García |first4=Nuria |last5=Laplana |first5=César |date=February 2020 |title=A cave lion (Panthera spelaea) skeleton from Torca del León (NW Iberia): Micromammals indicate a temperate and forest environment corresponding to GI-11 (MIS 3) |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119306420 |journal=Quaternary Science Reviews |volume=229 |pages=106123 |doi=10.1016/j.quascirev.2019.106123|bibcode=2020QSRv..22906123A |hdl=10651/54312 |hdl-access=free }}</ref> such as in Europe during the [[Last Interglacial|Last Interglacial/Eemian]].<ref>{{Cite journal |last=Diedrich |first=Cajus G. |date=June 2011 |title=A diseased Panthera leo spelaea (Goldfuss 1810) lioness from a forest elephant graveyard in the Late Pleistocene (Eemian) interglacial lake at Neumark-Nord, central Germany |url=http://www.tandfonline.com/doi/abs/10.1080/08912963.2010.507814 |journal=Historical Biology |volume=23 |issue=2–3 |pages=195–217 |doi=10.1080/08912963.2010.507814 |bibcode=2011HBio...23..195D |issn=0891-2963}}</ref>
+==Paleobiology==
-The cave lion probably inhabited predominantly open habitats such as [[steppe]] and grasslands although it would have also have occurred in open woodlands as well.<ref name=Stuart2011/> It also lived in mountainous regions, likely because it sought out hibernating bears in montane caves as a food source during the winter.<ref>{{Cite journal |last1=Sabol |first1=M. |last2=Gullár |first2=J. |last3=Horvát |first3=J. |date=2018 |title=Montane record of the late Pleistocene ''Panthera spelaea'' (Goldfuss, 1810) from the Západné Tatry Mountains (northern Slovakia) |journal=[[Journal of Vertebrate Paleontology]] |volume=38 |issue=3 |pages=e1467921 |doi=10.1080/02724634.2018.1467921 |doi-access=free|bibcode=2018JVPal..38E7921S }}</ref> While during the Last Glacial Period it was often associated with cold environments, the species also inhabited temperate environments,<ref>{{Cite journal |last1=Álvarez-Lao |first1=Diego J. |last2=Álvarez-Vena |first2=Adrián |last3=Ballesteros |first3=Daniel |last4=García |first4=Nuria |last5=Laplana |first5=César |date=February 2020 |title=A cave lion (Panthera spelaea) skeleton from Torca del León (NW Iberia): Micromammals indicate a temperate and forest environment corresponding to GI-11 (MIS 3) |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119306420 |journal=Quaternary Science Reviews |language=en |volume=229 |pages=106123 |doi=10.1016/j.quascirev.2019.106123|bibcode=2020QSRv..22906123A |hdl=10651/54312 |hdl-access=free }}</ref> such as in Europe during the [[Last Interglacial|Last Interglacial/Eemian]].<ref>{{Cite journal |last=Diedrich |first=Cajus G. |date=June 2011 |title=A diseased Panthera leo spelaea (Goldfuss 1810) lioness from a forest elephant graveyard in the Late Pleistocene (Eemian) interglacial lake at Neumark-Nord, central Germany |url=http://www.tandfonline.com/doi/abs/10.1080/08912963.2010.507814 |journal=Historical Biology |language=en |volume=23 |issue=2–3 |pages=195–217 |doi=10.1080/08912963.2010.507814 |bibcode=2011HBio...23..195D |issn=0891-2963}}</ref>
+=== Ecology ===
-==Paleobiology==
-=== Habitat ===
 [[File:Hoehlenloewe CaveLion hharder.jpg|thumb|right|A cave lion with a reindeer, painting by [[Heinrich Harder]]<ref>{{cite book |author=Bölsche, W. |name-list-style=amp |author2=Harder, H. |year=1900 |title=Tiere der Urwelt. Serie III |location=Wandsbek-Hamburg |publisher=Verlag der Kakao-Compagnie Theodor Reichardt}}</ref>]]
-''P. spelaea'' inhabited colder environments such as [[mammoth steppe]] and [[boreal forest]]. It was one of the [[keystone species]] of the mammoth steppe, being one of the main [[apex predator]]s alongside the [[gray wolf]], [[cave hyena]] and [[brown bear]].<ref name="Bocherens2015">{{Cite journal |last=Bocherens |first=H. |date=2015 |title=Isotopic tracking of large carnivore palaeoecology in the mammoth steppe |url= |url-status= |journal=[[Quaternary Science Reviews]] |volume=117 |pages=42–71 |bibcode=2015QSRv..117...42B |doi=10.1016/j.quascirev.2015.03.018 |archive-url= |archive-date= |access-date=}}</ref> Large amounts of bones belonging to ''P. spelaea'' were excavated in caves, where bones of cave hyena, [[cave bear]] and [[Paleolithic]] artefacts were also found.<ref>{{cite journal |author=Diedrich, C. G. |year=2011 |title=The largest European lion ''Panthera leo spelaea'' (Goldfuss 1810) population from the Zoolithen Cave, Germany: specialised cave bear predators of Europe |journal=Historical Biology |volume=23 |issue=2–3 |pages=271–311 |doi=10.1080/08912963.2010.546529 |bibcode=2011HBio...23..271D |s2cid=86638786}}</ref><ref>{{cite journal |author=Diedrich, C. G. |year=2011 |title=Pleistocene ''Panthera leo spelaea'' (Goldfuss 1810) remains from the Balve cave (NW Germany) - a cave bear, hyena den and middle palaeolithic human cave – and review of the Sauerland Karst lion cave sites |journal=Quaternaire |volume=22 |issue=2 |pages=105–127 |doi=10.4000/quaternaire.5897 |url=http://journals.openedition.org/quaternaire/5897 |access-date=18 September 2020 |archive-date=11 April 2021 |archive-url=https://web.archive.org/web/20210411153241/https://journals.openedition.org/quaternaire/5897 |url-status=live }}</ref> Some of these accumulations of cave lion bones have been attributed to hoarding of meat from cave lion carcasses by cave hyenas in caves occupied by the latter.<ref>{{Cite journal |last=Diedrich |first=Cajus G. |date=1 July 2011 |title=Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss, 1810) footprints and bone records from open air sites in northern Germany – Evidence of hyena-lion antagonism and scavenging in Europe |url=https://www.sciencedirect.com/science/article/pii/S0277379111000783 |journal=[[Quaternary Science Reviews]] |volume=30 |issue=15 |pages=1883–1906 |doi=10.1016/j.quascirev.2011.03.006 |bibcode=2011QSRv...30.1883D |issn=0277-3791 |access-date=4 February 2024 |via=Elsevier Science Direct |archive-date=18 May 2024 |archive-url=https://web.archive.org/web/20240518071940/https://www.sciencedirect.com/science/article/abs/pii/S0277379111000783 |url-status=live }}</ref>
+''P. spelaea'' was one of the [[keystone species]] of the mammoth steppe, being one of the main [[apex predator]]s alongside the [[gray wolf]], [[cave hyena]] and [[brown bear]].<ref name="Bocherens2015">{{Cite journal |last=Bocherens |first=H. |date=2015 |title=Isotopic tracking of large carnivore palaeoecology in the mammoth steppe |url= |journal=[[Quaternary Science Reviews]] |volume=117 |pages=42–71 |bibcode=2015QSRv..117...42B |doi=10.1016/j.quascirev.2015.03.018 |access-date=}}</ref> Large amounts of bones belonging to ''P. spelaea'' were excavated in caves, where bones of cave hyena, [[cave bear]] and [[Paleolithic]] artefacts were also found.<ref>{{cite journal |author=Diedrich, C. G. |year=2011 |title=The largest European lion ''Panthera leo spelaea'' (Goldfuss 1810) population from the Zoolithen Cave, Germany: specialised cave bear predators of Europe |journal=Historical Biology |volume=23 |issue=2–3 |pages=271–311 |doi=10.1080/08912963.2010.546529 |bibcode=2011HBio...23..271D |s2cid=86638786}}</ref><ref>{{cite journal |author=Diedrich, C. G. |year=2011 |title=Pleistocene ''Panthera leo spelaea'' (Goldfuss 1810) remains from the Balve cave (NW Germany) - a cave bear, hyena den and middle palaeolithic human cave – and review of the Sauerland Karst lion cave sites |journal=Quaternaire |volume=22 |issue=2 |pages=105–127 |doi=10.4000/quaternaire.5897 |url=http://journals.openedition.org/quaternaire/5897 |access-date=18 September 2020 |archive-date=11 April 2021 |archive-url=https://web.archive.org/web/20210411153241/https://journals.openedition.org/quaternaire/5897 |url-status=live }}</ref> Despite their common name, "cave lions" probably only infrequently if ever used caves, and were present in regions where caves were absent.<ref name="diedrich&zak">Diedrich, C.G. & ŽÁK, K. 2006. [http://www.geology.cz/bulletin/fulltext/bullgeosci200604237.pdf Prey deposits and den sites of the Upper Pleistocene hyena Crocuta crocuta spelaea (Goldfuss, 1823) in horizontal and vertical caves of the Bohemian Karst (Czech Republic)]. ''Bulletin of Geosciences'' 81(4), 237–276 (25 figures). Czech Geological Survey, Prague. ISSN 1214-1119.</ref> Some of these accumulations of cave lion bones in cave hyena dens have been attributed to confrontations between cave hyenas and cave lions over carcasses, with the remains of cave lions killed in these confrontations subsequently transported to the dens.<ref>{{Cite journal |last=Diedrich |first=Cajus G. |date=1 July 2011 |title=Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss, 1810) footprints and bone records from open air sites in northern Germany – Evidence of hyena-lion antagonism and scavenging in Europe |url=https://www.sciencedirect.com/science/article/pii/S0277379111000783 |journal=[[Quaternary Science Reviews]] |volume=30 |issue=15 |pages=1883–1906 |doi=10.1016/j.quascirev.2011.03.006 |bibcode=2011QSRv...30.1883D |issn=0277-3791 |access-date=4 February 2024 |via=Elsevier Science Direct |archive-date=18 May 2024 |archive-url=https://web.archive.org/web/20240518071940/https://www.sciencedirect.com/science/article/abs/pii/S0277379111000783 |url-status=live }}</ref><ref name="diedrich&zak" />
-=== Social Behavior ===
-Whether or not cave lions existed in prides like modern lions is unclear. Isotopic analysis done by Hervé Bocherns in 2015, suggested cave lions in Europe may have been solitary, due to scattering of individual data which was more similar to individualistic behavior compared to modern day lion populations.<ref name="Bocherens2015" />
+Isotopic analyses of bone [[collagen]] samples extracted from remains in Europe<ref name="Bocherens" /><ref name="Bocherens2015" /> and East Beringia<ref>{{Cite journal |last=Drucker |first=Dorothée G. |date=2022-05-31 |title=The Isotopic Ecology of the Mammoth Steppe |url=https://www.annualreviews.org/doi/10.1146/annurev-earth-100821-081832 |journal=Annual Review of Earth and Planetary Sciences |language=en |volume=50 |issue=1 |pages=395–418 |doi=10.1146/annurev-earth-100821-081832 |issn=0084-6597}}</ref> indicate that [[reindeer]] were particularly prominent in the diet of cave lions in these regions during the Last Glacial Period.<ref name="Bocherens2015" /> Cave lions also seem to have opportunistically preyed on the cubs of [[Cave bear|cave bears]].<ref name="Bocherens2015" /><ref name="Bocherens">{{cite journal |author1=Bocherens, H. |author2=Drucker, D. G. |author3=Bonjean, D. |author4=Bridault, A. |author5=Conard, N. J. |author6=Cupillard, C. |author7=Germonpré, M. |author8=Höneisen, M. |author9=Münzel, S. C. |author10=Napierala, H. |author11=Patou-Mathis, M. |name-list-style=amp |date=2011 |title=Isotopic evidence for dietary ecology of cave lion (''Panthera spelaea'') in North-Western Europe: prey choice, competition and implications for extinction |url=https://hal.archives-ouvertes.fr/hal-01673488/file/BOCHERENS%20%26%20al%202011%28QI-lion%29.pdf |url-status=live |journal=Quaternary International |volume=245 |issue=2 |pages=249–261 |bibcode=2011QuInt.245..249B |doi=10.1016/j.quaint.2011.02.023 |s2cid=129706533 |archive-url=https://web.archive.org/web/20200511073456/https://hal.archives-ouvertes.fr/hal-01673488/file/BOCHERENS%20%26%20al%202011%28QI-lion%29.pdf |archive-date=11 May 2020 |access-date=18 September 2020}}</ref> Isotopic analysis of other European specimens suggests a diet including [[wild horse]], [[woolly mammoth]] and cave bears for these individuals.<ref name="Bocherens2015" />
-Boeskorov et al. 2021 suggested both European and Beringian cave lions could've  hunted in larger prides than modern lions because sexual dimorphism in the dentition of cave lions was larger than solitude felids and African lions. However, they admitted the data is insufficient to come down to a proper conclusion.<ref name="Boeskorov_al.2021" />{{Notetag|This analysis would contradict the scattering of individual data of European cave lions from Bocherens 2015. The same study mentioned cave hyenas were more dominant against cave lions when their range overlapped with one another, via cave lions mainly favoring reindeer, while cave hyena hunted all large herbivores that were available. Beringian cave lions had a more varied diet because cave hyenas aren't found in east Berigina.|group=note}}
+Other possible prey species were [[giant deer]], [[red deer]], [[muskox]], [[aurochs]], [[wisent]], [[bison priscus|steppe bison]], and young [[woolly rhinoceros]]. It likely competed for prey with the [[Panthera pardus spelaea|European leopards]], cave hyenas, brown bears and grey wolves in Eurasia,<ref name="Diedrich 2013">{{cite journal |author=Diedrich, C. G. |year=2013 |title=Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art |journal=Quaternary Science Reviews |volume=76 |pages=167–193 |bibcode=2013QSRv...76..167D |doi=10.1016/j.quascirev.2013.05.009}}</ref> along with [[Arctodus|short-faced bears]], ''[[Homotherium]]'', and [[Beringian wolf|Beringian wolves]] in [[Beringia]].<ref name="Bocherens2015" />
-=== Diet ===
-Isotopic analyses of bone [[collagen]] samples extracted from fossils indicate that cave bear cubs, [[reindeer]] and other [[cervid]]s (deer) were prominent in the diet of cave lions. Later cave lions seem to have preyed foremost on reindeer, up to the brink of local extinction or extirpation of both species.<ref name="Bocherens2015" /><ref name=Bocherens>{{cite journal |author1=Bocherens, H. |name-list-style=amp |author2=Drucker, D. G. |author3=Bonjean, D. |author4=Bridault, A. |author5=Conard, N. J. |author6=Cupillard, C. |author7=Germonpré, M. |author8=Höneisen, M. |author9=Münzel, S. C. |author10=Napierala, H. |author11=Patou-Mathis, M. |date=2011 |title=Isotopic evidence for dietary ecology of cave lion (''Panthera spelaea'') in North-Western Europe: prey choice, competition and implications for extinction |journal=Quaternary International |volume=245 |issue=2 |pages=249–261 |url=https://hal.archives-ouvertes.fr/hal-01673488/file/BOCHERENS%20%26%20al%202011%28QI-lion%29.pdf |doi=10.1016/j.quaint.2011.02.023 |bibcode=2011QuInt.245..249B |s2cid=129706533 |access-date=18 September 2020 |archive-date=11 May 2020 |archive-url=https://web.archive.org/web/20200511073456/https://hal.archives-ouvertes.fr/hal-01673488/file/BOCHERENS%20%26%20al%202011%28QI-lion%29.pdf |url-status=live }}</ref> Other possible prey species were [[giant deer]], [[red deer]], [[wild horse]], [[muskox]], [[aurochs]], [[wisent]], [[bison priscus|steppe bison]], young [[woolly rhino]], and young [[woolly mammoth]]. It likely competed for prey with the [[Panthera pardus spelaea|European Ice Age leopard]], cave hyenas, brown bears and grey wolves in Eurasia,<ref name="Diedrich 2013">{{cite journal |author=Diedrich, C. G. |year=2013 |title=Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art |journal=Quaternary Science Reviews |pages=167–193 |doi=10.1016/j.quascirev.2013.05.009 |volume=76|bibcode=2013QSRv...76..167D}}</ref> along with [[Arctodus|short-faced bears]], ''[[Homotherium]]'', and [[Beringian wolf|Beringian wolves]] in [[Beringia]].<ref name="Bocherens2015" />
+=== Social behavior ===
+Whether or not cave lions existed in prides like modern lions is unclear. Isotopic analysis done by Hervé Bocherns in 2015, suggested cave lions in Europe may have been solitary, due to scattering of individual data which was more similar to individualistic behavior compared to modern day lion populations.<ref name="Bocherens2015" /> Some other authors have also argued that the absence of manes in cave lions suggests that cave lions did not live in prides, given the importance of manes in the social hierarchy of modern lions.<ref name="Boeskorov_al.2021" /> Boeskorov et al. 2021 suggested both European and Beringian cave lions may have hunted in larger prides than modern lions because sexual dimorphism in cave lions was more pronounced than in modern African lions and solitary big cats. However, they admitted the data is insufficient to come down to a certain conclusion.<ref name="Boeskorov_al.2021" />
+Cave lion cubs appear to have lived in dens during their earliest stages of life, like modern lion cubs and were likely solely raised by females, like living ''Panthera'' species.<ref name="Boeskorov_al.2021" />
 == Relationship with humans ==
+{{multiple image
-There is direct evidence of hunting of cave lions and exploitation of their pelts in Europe by both [[Neanderthal|Neanderthals]] during the [[Middle Paleolithic]],<ref name=":3">{{Cite journal |last1=Russo |first1=Gabriele |last2=Milks |first2=Annemieke |last3=Leder |first3=Dirk |last4=Koddenberg |first4=Tim |last5=Starkovich |first5=Britt M. |last6=Duval |first6=M. |last7=Zhao |first7=J.-X. |last8=Darga |first8=Robert |last9=Rosendahl |first9=Wilfried |last10=Terberger |first10=Thomas |date=2023-10-12 |title=First direct evidence of lion hunting and the early use of a lion pelt by Neanderthals |journal=Scientific Reports |language=en |volume=13 |issue=1 |page=16405 |doi=10.1038/s41598-023-42764-0 |issn=2045-2322 |pmc=10570355 |pmid=37828055|bibcode=2023NatSR..1316405R }}</ref> as well as modern humans during [[Upper Paleolithic]] (as evidenced by the [[Magdalenian]] aged La Garma site in Spain).<ref name="UnderTheSkinOfLion">{{Cite journal |last1=Cueto |first1=Marián |last2=Camarós |first2=Edgard |last3=Castaños |first3=Pedro |last4=Ontañón |first4=Roberto |last5=Arias |first5=Pablo |date=26 October 2016 |editor-last=Macchiarelli |editor-first=Roberto |title=Under the Skin of a Lion: Unique Evidence of Upper Paleolithic Exploitation and Use of Cave Lion (Panthera spelaea) from the Lower Gallery of La Garma (Spain) |journal=[[PLOS ONE]] |language=en |volume=11 |issue=10 |pages=e0163591 |bibcode=2016PLoSO..1163591C |doi=10.1371/journal.pone.0163591 |issn=1932-6203 |pmc=5082676 |pmid=27783697 |doi-access=free}}</ref> Modern humans also drew cave paintings of cave lions, engraved their likeness on bones and created sculptures of them, including the famous anthropomorphic [[Lion-man|lion man]] figure from Germany dating to around 41-35,000 years ago with the body of a human and the head of a lion. Cave lion canines with perforated holes may have been worn as personal oraments.<ref name=":3" />{{gallery|Landesmuseum Württemberg -Vogelherdhöhle-Löwenköpfchen536.jpg|Carving of a ''Panthera spelaea'' head from [[Vogelherd Cave]], Germany, dating to around 35-40,000 years ago|Isturitz big cat.jpg|Drawing of a now-lost Paleolithic sculpture of a cave lion from Isturitz, France|GER — BW — RPT — Ulm — Marktpl. 9 — (Ulmer Museum) — Mattes 2022-03-13 Batch (34).jpg|The famous anthropomorphic [[Lion-man]] sculpture from [[Hohlenstein-Stadel]] cave, Germany|||||||width=185|height=|lines=|align=center|title=Paleolithic artwork of ''Panthera spelaea''}}
+| perrow            = 2
+| image1            = Isturitz big cat.jpg
+| caption1          = Drawing of a now-lost Palaeolithic sculpture of a cave lion from Isturitz, France
+| image2            = Lascaux-diverticule-félins.jpg
+| caption2          = Drawing of cave lions in the Chamber of Felines, Lascaux caves
+| image3            = Landesmuseum Württemberg -Vogelherdhöhle-Löwenköpfchen536.jpg
+| caption3          = Carving of a ''Panthera spelaea'' head from [[Vogelherd Cave]], dating to around 35–40,000 years ago
+| image4            = GER — BW — RPT — Ulm — Marktpl. 9 — (Ulmer Museum) — Mattes 2022-03-13 Batch (34).jpg
+| caption4          = Anthropomorphic [[Lion-man]] (Löwenmensch) figurine, around 35-41,000 years old
+}}
+Cave lions were hunted and their pelts exploited in Europe by [[Neanderthal]]s during the [[Middle Paleolithic]],<ref name=Russo>{{Cite journal |last1=Russo |first1=G. |last2=Milks |first2=A. |last3=Leder |first3=D.|last4=Koddenberg |first4=T. |last5=Starkovich |first5=B. M. |last6=Duval |first6=M. |last7=Zhao |first7=J.-X. |last8=Darga |first8=R. |last9=Rosendahl |first9=W. |last10=Terberger |first10=T. |date=2023 |title=First direct evidence of lion hunting and the early use of a lion pelt by Neanderthals |journal=Scientific Reports |volume=13 |issue=1 |page=16405 |doi=10.1038/s41598-023-42764-0 |pmc=10570355 |pmid=37828055 |bibcode=2023NatSR..1316405R}}</ref> and during the [[Upper Paleolithic]] by modern humans in Spain as evidenced in the La Garma site dating to the [[Magdalenian]].<ref name=UnderTheSkinOfLion>{{Cite journal |last1=Cueto |first1=M. |last2=Camarós |first2=E. |last3=Castaños |first3=P. |last4=Ontañón |first4=R. |last5=Arias |first5=P. |date=2016 |title=Under the skin of a Lion: Unique evidence of Upper Paleolithic exploitation and use of Cave Lion (P''anthera spelaea'') from the Lower Gallery of La Garma (Spain) |journal=[[PLOS ONE]] |volume=11 |issue=10 |page=e0163591 |bibcode=2016PLoSO..1163591C |doi=10.1371/journal.pone.0163591 |pmc=5082676 |pmid=27783697 |doi-access=free}}</ref> Modern humans also drew cave paintings of cave lions, engraved their likeness on bones and created sculptures of them, including the famous anthropomorphic [[lion-man]] figure from [[Hohlenstein-Stadel]] cave in Germany dating to around 41-35,000 years ago with the body of a human and the head of a lion. Cave lion canines with perforated holes may have been worn as personal ornaments.<ref name=Russo/> Decorated stones with engravings representing cave lions have been found in southern Italy.<ref>{{Cite journal |last=Sigari |first=D. |last2=Bourdier |first2=C. |last3=Conti |first3=C. |last4=Conti |first4=J. |last5=Forti |first5=L. |last6=García-Diez |first6=M. |last7=Lai |first7=G. |last8=Mazzini |first8=I. |last9=Pieruccini |first9=P. |last10=Sardella |first10=R. |date=15 June 2024 |title=The last cave lion of the late Upper Palaeolithic: The engraved feline of Grotta Romanelli (southern Italy) |journal=[[Quaternary Science Reviews]] |volume=334 |pages=108670 |doi=10.1016/j.quascirev.2024.108670}}</ref>
 == Extinction ==
+{{See also|Late Pleistocene extinctions}}
-Radiocarbon dating suggests that the species went extinct approximately simultaneously across its range during the last few thousand years of the Late Pleistocene, around 14-15,000 years ago, possibly surviving around 1000 years later in the far east North American portion of its range. This timing roughly corresponds to the onset of the [[Bølling–Allerød Interstadial]] warm period and the consequent collapse of the [[mammoth steppe]] ecosystem. The precise cause of its extinction is unclear, but possibly involved environmental change from open habitats to closed forests, changes in prey abundance, as well as human impact, though it is difficult to distentangle the precise causes of its extinction.<ref name="Stuart2011" /> Cave lions appear to have undergone a [[population bottleneck]] that considerably reduced their genetic diversity between 47,000 and 18,000 years ago, probably driven at least in part by climatic instability.<ref>{{Cite journal |last=Ersmark |first=Erik |last2=Orlando |first2=Ludovic |last3=Sandoval-Castellanos |first3=Edson |last4=Barnes |first4=Ian |last5=Barnett |first5=Ross |last6=Stuart |first6=Anthony |last7=Lister |first7=Adrian |last8=Dalén |first8=Love |date=2015-03-09 |title=Population Demography and Genetic Diversity in the Pleistocene Cave Lion |url=https://openquaternary.com/articles/10.5334/oq.aa |journal=Open Quaternary |language=en-US |volume=1 |issue=1 |doi=10.5334/oq.aa |issn=2055-298X}}</ref>
+Radiocarbon dating suggests that the species went extinct approximately simultaneously across its range during the last few thousand years of the Late Pleistocene, around 14-15,000 years ago, possibly surviving around 1000 years later in the far east North American portion of its range. This timing roughly corresponds to the onset of the [[Bølling–Allerød Interstadial]] warm period and the consequent collapse of the [[mammoth steppe]] ecosystem. The precise cause of its extinction is unclear, but possibly involved environmental change from open habitats to closed forests, changes in prey abundance, as well as human impact, though it is difficult to distentangle the precise causes of its extinction.<ref name="Stuart2011" /> Cave lions appear to have undergone a [[population bottleneck]] that considerably reduced their genetic diversity between 47,000 and 18,000 years ago, probably driven at least in part by climatic instability.<ref>{{Cite journal |last1=Ersmark |first1=Erik |last2=Orlando |first2=Ludovic |last3=Sandoval-Castellanos |first3=Edson |last4=Barnes |first4=Ian |last5=Barnett |first5=Ross |last6=Stuart |first6=Anthony |last7=Lister |first7=Adrian |last8=Dalén |first8=Love |date=2015-03-09 |title=Population Demography and Genetic Diversity in the Pleistocene Cave Lion |journal=Open Quaternary |volume=1 |issue=1 |page=4 |doi=10.5334/oq.aa |doi-access=free |issn=2055-298X}}</ref>
 == Mummified specimens ==
@@ Line 96: / Line 109: @@
 Research results indicate that the cubs were likely barely a week old at the time of their deaths, as their milk teeth had not fully erupted. Further evidence suggests the cubs were hidden at a den site until they were strong enough to follow their mother back to the pride, as with modern lions. Researchers believe that the cubs were trapped and killed by a landslide, and that the absence of oxygen underground hindered their decomposition and allowed the cubs to be preserved in such good condition. A second expedition to the site where the cubs were found was planned for 2016, in hopes of finding either the remains of a third cub or possibly the cubs' mother.<ref>{{cite news |author=<!-- Staff --> |date=2015 |title=Whiskers still bristling after more than 12,000 years in the Siberian cold |url=http://siberiantimes.com/science/casestudy/news/n0491-whiskers-still-bristling-after-more-than-10000-years-in-the-siberian-cold |url-status=live |archive-url=https://web.archive.org/web/20161216074651/http://siberiantimes.com/science/casestudy/news/n0491-whiskers-still-bristling-after-more-than-10000-years-in-the-siberian-cold/ |archive-date=16 December 2016 |access-date=25 November 2016 |work=The Siberian Times}}</ref>
-In 2017, another frozen specimen, thought to be a lion cub, was found in Yakutia on the banks of the Tirekhtyakh River ({{lang-ru|Тирехтях}}), a tributary of the [[Indigirka]] River. This male cub was thought to be slightly older than the 2015 cubs at the time of its death; it is estimated to have been around one and a half to two months.<ref>{{cite news |author=<!-- Staff --> |date=2017 |title=Extinct cave lion cub in 'perfect' condition found in Siberia rising cloning hopes |url=http://siberiantimes.com/science/casestudy/news/extinct-cave-lion-cub-in-perfect-condition-found-in-siberia-rising-cloning-hopes/ |url-status=live |archive-url=https://web.archive.org/web/20171112012143/http://siberiantimes.com/science/casestudy/news/extinct-cave-lion-cub-in-perfect-condition-found-in-siberia-rising-cloning-hopes/ |archive-date=12 November 2017 |access-date=12 November 2017 |work=The Siberian Times}}</ref> In 2018, another preserved carcass of a cub was found in a location {{cvt|15|m|ft|sigfig=1}} away. It was considered to be around a month old when it died approximately 50,000 years ago, and presumed to be a sibling of the male cub.<ref>{{cite news |last=Gertcyk |first=O. |date=2018 |title=Cute first pictures of new 50,000 year old cave lion cub found perfectly preserved in permafrost |url=https://siberiantimes.com/science/casestudy/news/cute-first-pictures-of-new-50000-year-old-cave-lion-cub-found-perfectly-preserved-in-permafrost-of-yakutia/ |url-status=live |archive-url=https://web.archive.org/web/20180914132515/https://siberiantimes.com/science/casestudy/news/cute-first-pictures-of-new-50000-year-old-cave-lion-cub-found-perfectly-preserved-in-permafrost-of-yakutia/ |archive-date=14 September 2018 |access-date=16 September 2018 |publisher=The Siberian Times}}</ref> However, [[carbon date|carbon dating]] showed them to have lived about 15,000 years apart, with the female estimated to have lived 28,000 years ago, and the male 43,448 years ago.<ref name="Boeskorov_al.2021" /> Both cubs were well preserved, albeit with a few damages, with the female possibly being the "best preserved" animal discovered from the [[Ice age]].<ref>{{cite news |last=Gertcyk |first=O. |date=9 August 2021 |title=Meet Sparta, the 'best preserved ice age animal ever found' |url=https://siberiantimes.com/other/others/news/meet-sparta-the-best-preserved-ice-age-animal-ever-found/ |url-status=live |archive-url=https://web.archive.org/web/20210812082545/https://siberiantimes.com/other/others/news/meet-sparta-the-best-preserved-ice-age-animal-ever-found/ |archive-date=12 August 2021 |access-date=13 August 2021 |newspaper=[[The Siberian Times]]}}</ref>
+In 2017, another frozen specimen, thought to be a lion cub, was found in Yakutia on the banks of the Tirekhtyakh River ({{langx|ru|Тирехтях}}), a tributary of the [[Indigirka]] River. This male cub was thought to be slightly older than the 2015 cubs at the time of its death; it is estimated to have been around one and a half to two months.<ref>{{cite news |author=<!-- Staff --> |date=2017 |title=Extinct cave lion cub in 'perfect' condition found in Siberia rising cloning hopes |url=http://siberiantimes.com/science/casestudy/news/extinct-cave-lion-cub-in-perfect-condition-found-in-siberia-rising-cloning-hopes/ |url-status=live |archive-url=https://web.archive.org/web/20171112012143/http://siberiantimes.com/science/casestudy/news/extinct-cave-lion-cub-in-perfect-condition-found-in-siberia-rising-cloning-hopes/ |archive-date=12 November 2017 |access-date=12 November 2017 |work=The Siberian Times}}</ref> In 2018, another preserved carcass of a cub was found in a location {{cvt|15|m|ft|sigfig=1}} away. It was considered to be around a month old when it died approximately 50,000 years ago, and presumed to be a sibling of the male cub.<ref>{{cite news |last=Gertcyk |first=O. |date=2018 |title=Cute first pictures of new 50,000 year old cave lion cub found perfectly preserved in permafrost |url=https://siberiantimes.com/science/casestudy/news/cute-first-pictures-of-new-50000-year-old-cave-lion-cub-found-perfectly-preserved-in-permafrost-of-yakutia/ |url-status=live |archive-url=https://web.archive.org/web/20180914132515/https://siberiantimes.com/science/casestudy/news/cute-first-pictures-of-new-50000-year-old-cave-lion-cub-found-perfectly-preserved-in-permafrost-of-yakutia/ |archive-date=14 September 2018 |access-date=16 September 2018 |publisher=The Siberian Times}}</ref> However, [[carbon date|carbon dating]] showed them to have lived about 15,000 years apart, with the female estimated to have lived 28,000 years ago, and the male 43,448 years ago.<ref name="Boeskorov_al.2021" /> Both cubs were well preserved, albeit with a few damages, with the female possibly being the "best preserved" animal discovered from the [[Ice age]].<ref>{{cite news |last=Gertcyk |first=O. |date=9 August 2021 |title=Meet Sparta, the 'best preserved ice age animal ever found' |url=https://siberiantimes.com/other/others/news/meet-sparta-the-best-preserved-ice-age-animal-ever-found/ |url-status=live |archive-url=https://web.archive.org/web/20210812082545/https://siberiantimes.com/other/others/news/meet-sparta-the-best-preserved-ice-age-animal-ever-found/ |archive-date=12 August 2021 |access-date=13 August 2021 |newspaper=[[The Siberian Times]]}}</ref>
 ==See also==
@@ Line 137: / Line 150: @@
 [[Category:Taxa named by Georg August Goldfuss]]
 [[Category:Apex predators]]
+[[Category:Species endangered by climate change]]